Biochemical composition changes can be linked to the tolerance of four grassland species under more persistent precipitation regimes.

Climate models suggest that the persistence of summer precipitation regimes (PRs) is on the rise, characterized by both longer dry and longer wet durations. These PR changes may alter plant biochemical composition and thereby their economic and ecological characteristics. However, impacts of PR persistence have primarily been studied at the community level, largely ignoring the biochemistry of individual species. Here, we analyzed biochemical components of four grassland species with varying sensitivity to PR persistence (Holcus lanatus, Phleum pratense, Lychnis flos-cuculi, Plantago lanceolata) along a range of increasingly persistent PRs (longer consecutive dry and wet periods) in a mesocosm experiment. The more persistent PRs decreased nonstructural sugars, whereas they increased lignin in all species, possibly reducing plant quality. The most sensitive species Lychnis seemed less capable of altering its biochemical composition in response to altered PRs, which may partly explain its higher sensitivity. The more tolerant species may have a more robust and dynamic biochemical network, which buffers the effects of changes in individual biochemical components on biomass. We conclude that the biochemical composition changes are important determinants for plant performance under increasingly persistent precipitation regimes.

Risk of short-term biodiversity loss under more persistent precipitation regimes.

Recent findings indicate that atmospheric warming increases the persistence of weather patterns in the mid-latitudes, resulting in sequences of longer dry and wet periods compared to historic averages. The alternation of progressively longer dry and wet extremes could increasingly select for species with a broad environmental tolerance. As a consequence, biodiversity may decline. Here we explore the relationship between the persistence of summer precipitation regimes and plant diversity by subjecting experimental grassland mesocosms to a gradient of dry-wet alternation frequencies whilst keeping the total precipitation constant. The gradient varied the duration of consecutive wet and dry periods, from 1 up to 60 days with or without precipitation, over a total of 120 days. An alternation of longer dry and wet spells led to a severe loss of species richness (up to -75% relative to the current rainfall pattern in W-Europe) and functional diversity (enhanced dominance of grasses relative to nitrogen (N)-fixers and non-N-fixing forbs). Loss of N-fixers and non-N-fixing forbs in severe treatments was linked to lower baseline competitive success and higher physiological sensitivity to changes in soil moisture compared to grasses. The extent of diversity losses also strongly depended on the timing of the dry and wet periods. Regimes in which long droughts (≥20 days) coincided with above-average temperatures showed significantly more physiological plant stress over the experimental period, greater plant mortality, and impoverished communities by the end of the season. Across all regimes, the duration of the longest period below permanent wilting point was an accurate predictor of mortality across the communities, indicating that increasingly persistent precipitation regimes may reduce opportunities for drought stress alleviation. We conclude that without recruitment, which was precluded in this experiment, summer precipitation regimes with longer dry and wet spells will likely diminish plant diversity, at least in the short term.

Pushing precipitation to the extremes in distributed experiments: recommendations for simulating wet and dry years.

Intensification of the global hydrological cycle, ranging from larger individual precipitation events to more extreme multiyear droughts, has the potential to cause widespread alterations in ecosystem structure and function. With evidence that the incidence of extreme precipitation years (defined statistically from historical precipitation records) is increasing, there is a clear need to identify ecosystems that are most vulnerable to these changes and understand why some ecosystems are more sensitive to extremes than others. To date, opportunistic studies of naturally occurring extreme precipitation years, combined with results from a relatively small number of experiments, have provided limited mechanistic understanding of differences in ecosystem sensitivity, suggesting that new approaches are needed. Coordinated distributed experiments (CDEs) arrayed across multiple ecosystem types and focused on water can enhance our understanding of differential ecosystem sensitivity to precipitation extremes, but there are many design challenges to overcome (e.g., cost, comparability, standardization). Here, we evaluate contemporary experimental approaches for manipulating precipitation under field conditions to inform the design of 'Drought-Net', a relatively low-cost CDE that simulates extreme precipitation years. A common method for imposing both dry and wet years is to alter each ambient precipitation event. We endorse this approach for imposing extreme precipitation years because it simultaneously alters other precipitation characteristics (i.e., event size) consistent with natural precipitation patterns. However, we do not advocate applying identical treatment levels at all sites - a common approach to standardization in CDEs. This is because precipitation variability varies >fivefold globally resulting in a wide range of ecosystem-specific thresholds for defining extreme precipitation years. For CDEs focused on precipitation extremes, treatments should be based on each site's past climatic characteristics. This approach, though not often used by ecologists, allows ecological responses to be directly compared across disparate ecosystems and climates, facilitating process-level understanding of ecosystem sensitivity to precipitation extremes.

Effects of nitrogen and water addition on trace element stoichiometry in five grassland species.

A 9-year manipulative experiment with nitrogen (N) and water addition, simulating increasing N deposition and changing precipitation regime, was conducted to investigate the bioavailability of trace elements, iron (Fe), manganese (Mn), copper (Cu), and zinc (Zn) in soil, and their uptake by plants under the two environmental change factors in a semi-arid grassland of Inner Mongolia. We measured concentrations of trace elements in soil and in foliage of five common herbaceous species including 3 forbs and 2 grasses. In addition, bioaccumulation factors (BAF, the ratio of the chemical concentration in the organism and the chemical concentration in the growth substrate) and foliar Fe:Mn ratio in each plant was calculated. Our results showed that soil available Fe, Mn and Cu concentrations increased under N addition and were negatively correlated with both soil pH and cation exchange capacity. Water addition partly counteracted the positive effects of N addition on available trace element concentrations in the soil. Foliar Mn, Cu and Zn concentrations increased but Fe concentration decreased with N addition, resulting in foliar elemental imbalances among Fe and other selected trace elements. Water addition alleviated the effect of N addition. Forbs are more likely to suffer from Mn toxicity and Fe deficiency than grass species, indicating more sensitivity to changing elemental bioavailability in soil. Our results suggested that soil acidification due to N deposition may accelerate trace element cycling and lead to elemental imbalance in soil-plant systems of semi-arid grasslands and these impacts of N deposition on semi-arid grasslands were affected by water addition. These findings indicate an important role for soil trace elements in maintaining ecosystem functions associated with atmospheric N deposition and changing precipitation regimes in the future.

Response of soil CO2 efflux to precipitation manipulation in a semiarid grassland.

Soil CO2 efflux (SCE) is an important component of ecosystem CO2 exchange and is largely temperature and moisture dependent, providing feedback between C cycling and the climate system. We used a precipitation manipulation experiment to examine the effects of precipitation treatment on SCE and its dependences on soil temperature and moisture in a semiarid grassland. Precipitation manipulation included ambient precipitation, decreased precipitation (-43%), or increased precipitation (+17%). The SCE was measured from July 2013 to December 2014, and CO2 emission during the experimental period was assessed. The response curves of SCE to soil temperature and moisture were analyzed to determine whether the dependence of SCE on soil temperature or moisture varied with precipitation manipulation. The SCE significantly varied seasonally but was not affected by precipitation treatments regardless of season. Increasing precipitation resulted in an upward shift of SCE-temperature response curves and rightward shift of SCE-moisture response curves, while decreasing precipitation resulted in opposite shifts of such response curves. These shifts in the SCE response curves suggested that increasing precipitation strengthened the dependence of SCE on temperature or moisture, and decreasing precipitation weakened such dependences. Such shifts affected the predictions in soil CO2 emissions for different precipitation treatments. When considering such shifts, decreasing or increasing precipitation resulted in 43 or 75% less change, respectively, in CO2 emission compared with changes in emissions predicted without considering such shifts. Furthermore, the effects of shifts in SCE response curves on CO2 emission prediction were greater during the growing than the non-growing season.

Zoning of precipitation regimes on the Qinghai-Tibet Plateau and its surrounding areas responded by the vegetation distribution.

Compared with other factors influencing vegetation patterns, such as light and temperature, precipitation has relatively large variability, especially on the Qinghai-Tibet Plateau (QTP), where the natural environment is extremely fragile and sensitive. However, the impact of precipitation regimes, rather than precipitation amount, on vegetation has seldom been revealed. This study characterised the precipitation regimes by both the amount and temporal distribution of precipitation and zoned the QTP as different precipitation regimes accordingly. The response of vegetation to such precipitation regimes was then investigated. The results indicate that the vegetation patterns are quite consistent with zoning, that is, there is a certain type or a few dominant types of vegetation in each sub-region divided by the precipitation regimes. The areas where the precipitation became more uniform within a year were concentrated in grassland and bare land, which benefits the restoration and improvement of the ecological environment of the plateau. The increase in precipitation variability in the south-eastern part of the plateau may lead to natural disasters such as floods and mudslides. This study provides a novel perspective to understand the distribution of vegetation patterns.

Responses of soil respiration to rainfall addition in a desert ecosystem: Linking physiological activities and rainfall pattern.

The tight linkage between photosynthesis (An) and soil respiration (Rs) has been verified in many terrestrial ecosystems. However, it remains unclear whether this linkage occurs in desert ecosystems, where water is considered an important trigger of carbon cycling. A field experiment was performed under seven simulated rainfall amounts (0, 3, 5, 10, 15, 25, and 40 mm) with two co-existing desert plants (Reaumuria soongorica and Nitraria sphaerocarpa) in June (early growing season, EGS) and August (middle growing season, MGS) in 2016. An, Rs, predawn water potential (Ψpd), soil temperature (Ts) and soil moisture (Swc) were measured for each treatment or control plot for 3 weeks. Our objective was to examine the effects of rainfall pattern on Rs and physiological responses of the two plants and the relationships between Rs and biotic and abiotic factors. No obvious variations in Ψpd or An were found under small rainfall events. However, when the rainfall amount exceeded 10 mm, both plants responded strongly, and the response patterns of Rs showed trends similar to those of An, which varied between species and seasons. Moreover, rain additions of 3-40 mm significantly increased Rs, and the relative changes in Rs (ΔRs) of both species were much larger in the EGS than in the MGS. Importantly, abiotic factors may have controlled the variations in Rs under small rain events while An played a more important role in regulating the variations in Rs when the rainfall amount exceeded 10 mm for both species, suggest that the rainfall pattern-driven changes in Rs composition interact with physiological activity and abiotic factors to regulate the response of Rs to rainfall variability in desert ecosystems. Thus, climate change in the coming decades may lead to carbon sequestration by desert plants, which may cause desert ecosystems to act as carbon sinks.