Complementary belowground strategies underlie species coexistence in an early successional forest.

Unravelling belowground strategies is critical for understanding species coexistence and successional dynamics; yet, our knowledge of nutrient acquisition strategies of forest species at different successional stages remains limited. We measured morphological (diameter, specific root length, and root tissue density), architectural (branching ratio), physiological (ammonium, nitrate, and glycine uptake rates) root traits, and mycorrhizal colonisation rates of eight coexisting woody species in an early successional plantation forest in subtropical China. By incorporating physiological uptake efficiency, we revealed a bi-dimensional root economics space comprising of an 'amount-efficiency' dimension represented by morphological and physiological traits, and a 'self-symbiosis' dimension dominated by architectural and mycorrhizal traits. The early pioneer species relied on root-fungal symbiosis, developing densely branched roots with high mycorrhizal colonisation rates for foraging mobile soil nitrate. The late pioneer species invested in roots themselves and allocated effort towards improving uptake efficiency of less-mobile ammonium. Within the root economics space, the covariation of axes with soil phosphorus availability also distinguished the strategy preference of the two successional groups. These results demonstrate the importance of incorporating physiological uptake efficiency into root economics space, and reveal a trade-off between expanding soil physical space exploration and improving physiological uptake efficiency for successional species coexistence in forests.

Mycorrhizal symbiosis pathway and edaphic fertility frame root economics space among tree species.

The root economics space (RES) is multidimensional and largely shaped by belowground biotic and abiotic influences. However, how root-fungal symbioses and edaphic fertility drive this complexity remains unclear. Here, we measured absorptive root traits of 112 tree species in temperate and subtropical forests of China, including traits linked to functional differences between arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) hosts. Our data, from known mycorrhizal tree species, revealed a 'fungal-symbiosis' dimension distinguishing AM from ECM species. This divergence likely resulted from the contrasting mycorrhizal evolutionary development of AM vs ECM associations. Increased root tissue cortical space facilitates AM symbiosis, whereas increased root branching favours ECM symbiosis. Irrespective of mycorrhizal type, a 'root-lifespan' dimension reflecting aspects of root construction cost and defence was controlled by variation in specific root length and root tissue density, which was fully independent of root nitrogen content. Within this function-based RES, we observed a substantial covariation of axes with soil phosphorus and nitrate levels, highlighting the role played by these two axes in nutrient acquisition and conservation. Overall, our findings demonstrate the importance of evolved mycorrhizal symbiosis pathway and edaphic fertility in framing the RES, and provide theoretical and mechanistic insights into the complexity of root economics.

Assessment of vegetation change on the Mongolian Plateau over three decades using different remote sensing products.

As a major component of temperate steppes in the Eurasian continent, the Mongolian Plateau (MP) plays a pivotal role in the East Asian and global carbon cycles. This paper describes the use of five remote sensing indices derived from satellite data to characterize vegetation cover on MP, namely: gross primary production (GPP), net primary production (NPP), normalized difference vegetation index (NDVI), leaf area index (LAI) and fractional vegetation cover (FVC). It is found that GPP, NPP, and NDVI exhibit increasing trends, whereas LAI and FVC present decreasing trends on the MP since 1982. The different indices highlight discrepancies in the spatial pattern of vegetation growth, with the greatest increase in the southeast of MP. Only 3.4% of the total land area of MP exhibited consistent trends in the indices (0.1% degradation and 3.3% growth, P < 0.01), with the synchronous change of both LAI and NPP exhibiting higher consistency than that of raw NDVI and NPP. Understanding of the characteristics and status of vegetation change on the MP has far-reaching implications for its ecological protection management, and climate change mitigation.

Mycorrhizal and environmental controls over root trait-decomposition linkage of woody trees.

Traits are critical in predicting decomposition that fuels carbon and nutrient cycling in ecosystems. However, our understanding of root trait-decomposition linkage, and especially its dependence on mycorrhizal type and environmental context, remains limited. We explored the control of morphological and chemical (carbon- and nutrient-related) traits over decomposition of absorptive roots in 30 tree species associated with either arbuscular mycorrhizal (AM) or ectomycorrhizal (ECM) fungi in temperate and subtropical forests in China. Carbon-related traits (acid-unhydrolysable residue (AUR) and cellulose concentrations) had predominant control of root decomposition in AM species while nutrient-related traits (magnesium concentration) predominately controlled that in ECM species. Thicker absorptive roots decomposed faster in AM species as a result of their lower AUR concentrations, but more slowly in ECM angiosperm species potentially as a result of their higher magnesium concentrations. Root decomposition was linked to root nutrient economy in both forests while root diameter-decomposition coordination emerged only in the subtropical forest where root diameter and decomposition presented similar cross-species variations. Our findings suggest that root trait-decomposition linkages differ strongly with mycorrhizal type and environment, and that root diameter can predict decomposition but in opposing directions and with contrasting mechanisms for AM and ECM species.

Carbon pools in China's terrestrial ecosystems: New estimates based on an intensive field survey.

China's terrestrial ecosystems have functioned as important carbon sinks. However, previous estimates of carbon budgets have included large uncertainties owing to the limitations of sample size, multiple data sources, and inconsistent methodologies. In this study, we conducted an intensive field campaign involving 14,371 field plots to investigate all sectors of carbon stocks in China's forests, shrublands, grasslands, and croplands to better estimate the regional and national carbon pools and to explore the biogeographical patterns and potential drivers of these pools. The total carbon pool in these four ecosystems was 79.24 ± 2.42 Pg C, of which 82.9% was stored in soil (to a depth of 1 m), 16.5% in biomass, and 0.60% in litter. Forests, shrublands, grasslands, and croplands contained 30.83 ± 1.57 Pg C, 6.69 ± 0.32 Pg C, 25.40 ± 1.49 Pg C, and 16.32 ± 0.41 Pg C, respectively. When all terrestrial ecosystems are taken into account, the country's total carbon pool is 89.27 ± 1.05 Pg C. The carbon density of the forests, shrublands, and grasslands exhibited a strong correlation with climate: it decreased with increasing temperature but increased with increasing precipitation. Our analysis also suggests a significant sequestration potential of 1.9-3.4 Pg C in forest biomass in the next 10-20 years assuming no removals, mainly because of forest growth. Our results update the estimates of carbon pools in China's terrestrial ecosystems based on direct field measurements, and these estimates are essential to the validation and parameterization of carbon models in China and globally.

Unaltered phenology but increased production of ectomycorrhizal roots of Pinus elliottii under 4 years of nitrogen addition.

Timing (phenology) and amount (production) are two integral facets of root growth, and their shifts have profound influences on below-ground resource acquisition. However, the environmental control and the effects of nitrogen (N) deposition on the production and phenology of ectomycorrhizal (ECM) roots remain unclear. Using a 4 yr minirhizotron experiment, we explored the control of the production and three phenophases (initiation, peak, and cessation of growth) of ECM roots in two soil layers and investigated their dynamic responses to N addition in a seasonally dry subtropical Pinus elliottii forest. We found a stronger control of water availability on the production and a stronger control of temperature on the phenology of ECM roots under ambient conditions. Temperature was correlated positively with initiation and negatively with cessation, especially in the shallow layer. N addition did not affect the phenology of ECM roots but increased their production by modifying N and phosphorus (P) stoichiometry in the soil and foliage. Our findings suggest a greater sensitivity of production than phenology of ECM roots to N addition. The increased production of ECM roots under N addition could be driven by N-induced P limitation or some combination of below-ground resources (P, N, water).

Product and Metal Stocks Accumulation of China's Megacities: Patterns, Drivers, and Implications.

The rapid urbanization in China since the 1970s has led to an exponential growth of metal stocks (MS) in use in cities. A retrospect on the quantity, quality, and patterns of these MS is a prerequisite for projecting future metal demand, identifying urban mining potentials of metals, and informing sustainable urbanization strategies. Here, we deployed a bottom-up stock accounting method to estimate stocks of iron, copper, and aluminum embodied in 51 categories of products and infrastructure across 10 Chinese megacities from 1980 to 2016. We found that the MS in Chinese megacities had reached a level of 2.6-6.3 t/cap (on average 3.7 t/cap for iron, 58 kg/cap for copper, and 151 kg/cap for aluminum) in 2016, which still remained behind the level of western cities or potential saturation level on the country level (e.g., approximately 13 t/cap for iron). Economic development was identified as the most powerful driver for MS growth based on an IPAT decomposition analysis, indicating further increase in MS as China's urbanization and economic growth continues in the next decades. The latecomer cities should therefore explore a wide range of strategies, from urban planning to economy structure to regulations, for a transition toward more "metal-efficient" urbanization pathways.

Shifts in the dynamics of productivity signal ecosystem state transitions at the biome-scale.

Understanding ecosystem dynamics and predicting directional changes in ecosystem in response to global changes are ongoing challenges in ecology. Here we present a framework that links productivity dynamics and ecosystem state transitions based on a spatially continuous dataset of aboveground net primary productivity (ANPP) from the temperate grassland of China. Across a regional precipitation gradient, we quantified spatial patterns in ANPP dynamics (variability, asymmetry and sensitivity to rainfall) and related these to transitions from desert to semi-arid to mesic steppe. We show that these three indices of ANPP dynamics displayed distinct spatial patterns, with peaks signalling transitions between grassland types. Thus, monitoring shifts in ANPP dynamics has the potential for predicting ecosystem state transitions in the future. Current ecosystem models fail to capture these dynamics, highlighting the need to incorporate more nuanced ecological controls of productivity in models to forecast future ecosystem shifts.

Nitrogen deposition increases root production and turnover but slows root decomposition in Pinus elliottii plantations.

Fine roots of woody plants comprise multiple root orders, which can be functionally partitioned into two pools: absorptive fine roots (AFRs, orders 1, 2) and transport fine roots (TFRs, orders 3-5). However, the function-based fine-root dynamics and especially their responses to increased nitrogen (N) availability remain unclear. We explored dynamic responses of both AFRs and TFRs of Pinus elliottii to N addition in subtropical China based on a 4-yr minirhizotron experiment and a two-stage - early (0.5 yr) vs late (4 yr) - decomposition experiment. N addition increased the production, mortality, and turnover of AFRs but not TFRs. High rates of N persistently inhibited AFR decomposition but affected TFR decomposition differentially at the early (no effect) and late (negative effect) stages. The increased production of AFRs was driven by N-induced decrease in foliar and soil phosphorus (P) concentrations. The decreased decomposition of AFRs might be due to the increased acid-unhydrolyzable residues in decomposing roots. AFRs are the resource-acquiring module, the increased carbon allocation to AFRs may represent a P-acquiring strategy when N no longer limits growth of P. elliottii. Our results suggest that AFRs and TFRs respond differently to N deposition, both in terms of production, mortality, and turnover and in terms of decomposition.

A global synthesis of the rate and temperature sensitivity of soil nitrogen mineralization: latitudinal patterns and mechanisms.

Soil net nitrogen (N) mineralization (Nmin ) is a pivotal process in the global N cycle regulating the N availability of plant growth. Understanding the spatial patterns of Nmin, its temperature sensitivity (Q10 ) and regulatory mechanisms is critical for improving the management of soil nutrients. In this study, we evaluated 379 peer-reviewed scientific papers to explore how Nmin and the Q10 of Nmin varied among different ecosystems and regions at the global scale. The results showed that Nmin varied significantly among different ecosystems with a global average of 2.41 mg N soil kg-1  day-1 . Furthermore, Nmin significantly decreased with increasing latitude and altitude. The Q10 varied significantly among different ecosystems with a global average of 2.21, ranging from the highest found in forest soils (2.43) and the lowest found for grassland soils (1.67) and significantly increased with increasing latitude. Path analyses indicated that Nmin was primarily affected by the content of soil organic carbon (C), soil C:N ratio, and clay content, where Q10 was primarily influenced by the soil C:N ratio and soil pH. Furthermore, the activation energy (Ea ) of soil N mineralization was significantly and negative correlated with the substrate quality index among all ecosystems, indicating the applicability of the carbon quality temperature hypothesis to soil N mineralization at a global scale. These findings provided empirical evidence supporting that soil N availability, under global warming scenarios, is expected to increase stronger in colder regions as compared with that low-latitude regions due to the higher Q10 . This may alleviate the restriction of N supply for increased primary productivity at higher latitudes.

A synthesis of the effect of grazing exclusion on carbon dynamics in grasslands in China.

Grazing exclusion (GE) is considered to be an effective approach to restore degraded grasslands and to improve their carbon (C) sequestration. However, the C dynamics and related controlling factors in grasslands with GE have not been well characterized. This synthesis examines the dynamics of soil C content and vegetation biomass with the recovery age through synthesizing results of 51 sites in grasslands in China. The results illustrate increases in soil C content and vegetation biomass with GE at most sites. Generally, both soil C content and vegetation biomass arrive at steady state after 15 years of GE. In comparison, the rates of increase in above- and belowground biomass declined exponentially with the age of GE, whereas soil C content declined in a milder (linear) way, implying a lagged response of soil C to the inputs from plant biomass. Mean annual precipitation (MAP) and the rate of soil nitrogen (N) change were the main factors affecting the rate of soil C content change. MAP played a major role at the early stage, whereas the rate of soil N change was the major contributor at the middle and late stages. Our results imply that the national grassland restoration projects in China may be more beneficial for C sequestration in humid regions with high MAP. In addition, increased soil N supply to grasslands with GE at the latter recovery stage may enhance ecosystem C sequestration capacity.

Diverse responses of surface biogeophysical parameters to accelerated development and senescence of vegetation on the Mongolian Plateau.

Vegetation dynamics is essential for characterizing surface biogeophysical parameters. Speeds of vegetation development and senescence are well documented, however, the effects of vegetation growth rates on surface parameters during different growth stages remains unclear. By using such methods as trend analyses and correlation analyses, this study examines the variations and interactive relationships of leaf area index (LAI) and surface parameters including Albedo, evapotranspiration (ET), and land surface temperature (LST), derived from Moderate Resolution Imaging Spectroradiometer (MODIS), during the intra-growing season (April-October, GS) on the Mongolian Plateau (MP). Generally, LAI exhibited a significant upward trend across GS months. Significant changes in VLAI (the difference in LAI between 2 consecutive months) in April-May and September-October indicated that the vegetation change rates were accelerated in the early GS (April-June) and late GS (September-October). The effect of vegetation activity on surface parameters varies over time and space. The effects of VLAI on the speed of surface parameters were inconsistent during the intra-GS. As a result of the significant changes in LAI, VET (the difference in ET between 2 consecutive months) displayed a significant upward trend during the early GS but a significant downward trend during the late GS. With acceleration of vegetation activity, the effects of VET and VAlbedo (the difference in Albedo between 2 consecutive months) on LST could offset each other at different stages of the GS. In addition, the effect of VLAI on the speed of surface parameters varied significantly by vegetation types. Our findings imply that clarifying the impact of vegetation activity on surface parameters at different growth stages can advance our understanding of vegetation responses and feedbacks to climate change.

A trait-based root acquisition-defence-decomposition framework in angiosperm tree species.

To adapt to the complex belowground environment, plants make trade-offs between root resource acquisition and defence ability. This includes forming partnerships with different types of root associating microorganisms, such as arbuscular mycorrhizal and ectomycorrhizal fungi. These trade-offs, by mediating root chemistry, exert legacy effects on nutrient release during decomposition, which may, in turn, affect the ability of new roots to re-acquire resources, thereby generating a feedback loop. However, the linkages at the basis of this potential feedback loop remain largely unquantified. Here, we propose a trait-based root 'acquisition-defence-decomposition' conceptual framework and test the strength of relevant linkages across 90 angiosperm tree species. We show that, at the plant species level, the root-fungal symbiosis gradient within the root economics space, root chemical defence (condensed tannins), and root decomposition rate are closely linked, providing support to this framework. Beyond the dichotomy between arbuscular mycorrhizal-dominated versus ectomycorrhizal-dominated systems, we suggest a continuous shift in feedback loops, from 'high arbuscular mycorrhizal symbiosis-low defence-fast decomposition-inorganic nutrition' by evolutionarily ancient taxa to 'high ectomycorrhizal symbiosis-high defence-slow decomposition-organic nutrition' by more modern taxa. This 'acquisition-defence-decomposition' framework provides a foundation for testable hypotheses on multidimensional linkages between species' belowground strategies and ecosystem nutrient cycling in an evolutionary context.

Diverse responses of gross primary production and leaf area index to drought on the Mongolian Plateau.

Drought is a crucial factor regulating vegetation growth on the Mongolian Plateau (MP). Previous studies of drought effects on the MP have mainly concentrated on drought characterization, while the response of vegetation to drought remains unclear. To close this knowledge gap, we examined the response of MP vegetation to drought in terms of gross primary production (GPP) and leaf area index (LAI) from 1982 to 2018. Our findings show that intra-seasonally the frequency of drought occurrence in autumn had a greater impact on GPP (relative importance over 70 %), while the intensity of drought was more influential for LAI (relative importance approximately 60 %). Inter-seasonally, summer droughts had the most pronounced effect on vegetation (with median standardized anomalies of -0.72 for GPP and -0.4 for LAI, respectively). Additionally, we found that meteorological drought was more consistent with atmospheric aridity (high vapor pressure deficit) than soil drought (low soil moisture). This study advances knowledge of vegetation's susceptibility to climate extremes and improves the precision of predicting ecosystem response to climate change.

Plant-soil feedback regulates the trade-off between phosphorus acquisition pathways in Pinus elliottii.

Plant-soil feedback (PSF) is conventionally characterized by plant biomass growth, yet it remains unclear how PSF affects plant nutrient acquisition strategies (e.g., nutrient absorption and nutrient resorption) associated with plant growth, particularly under changing soil environments. A greenhouse experiment was performed with seedlings of Pinus elliottii Englem and conditioned soils of monoculture plantations (P. elliottii and Cunninghamia lanceolata Hook). Soil sterilization was designed to test plant phosphorus (P) acquisition strategy with and without native soil fungal communities. Soils from P. elliottii and C. lanceolata plantations were used to explore the specific soil legacy effects on two different P acquisition pathways (absorption and resorption). Phosphorus addition was also applied to examine the separate and combined effects of soil abiotic factors and soil fungal factors on P acquisition pathways. Due to diminished mycorrhizal symbiosis, PSF prompted plants to increasingly rely on P resorption under soil sterilization. In contrast, P absorption was employed preferentially in the heterospecific soil, where species-specific pathogenic fungi could not affect P absorption. Higher soil P availability diluted the effects of soil fungal factors on the trade-off between the two P acquisition pathways in terms of the absolute PSF. Moreover, P addition plays a limited role in terms of the relative PSF and does not affect the direction and strength of relative PSF. Our results reveal the role of PSF in regulating plant P acquisition pathways and highlight the interaction between mycorrhizal and pathogenic fungi as the underlying mechanism of PSF.

Predicting ecosystem productivity based on plant community traits.

With the rapid accumulation of plant trait data, major opportunities have arisen for the integration of these data into predicting ecosystem primary productivity across a range of spatial extents. Traditionally, traits have been used to explain physiological productivity at cell, organ, or plant scales, but scaling up to the ecosystem scale has remained challenging. Here, we show the need to combine measures of community-level traits and environmental factors to predict ecosystem productivity at landscape or biogeographic scales. We show how theory can extend the production ecology equation to enormous potential for integrating traits into ecological models that estimate productivity-related ecosystem functions across ecological scales and to anticipate the response of terrestrial ecosystems to global change.

Dew water isotopic ratios and their relationships to ecosystem water pools and fluxes in a cropland and a grassland in China.

Dew formation has the potential to modulate the spatial and temporal variations of isotopic contents of atmospheric water vapor, oxygen and carbon dioxide. The goal of this paper is to improve our understanding of the isotopic interactions between dew water and ecosystem water pools and fluxes through two field experiments in a wheat/maize cropland and in a short steppe grassland in China. Measurements were made during 94 dew events of the D and (18)O compositions of dew, atmospheric vapor, leaf, xylem and soil water, and the whole ecosystem water flux. Our results demonstrate that the equilibrium fractionation played a dominant role over the kinetic fractionation in controlling the dew water isotopic compositions. A significant correlation between the isotopic compositions of leaf water and dew water suggests a large role of top-down exchange with atmospheric vapor controlling the leaf water turnover at night. According to the isotopic labeling, dew water consisted of a downward flux of water vapor from above the canopy (98%) and upward fluxes originated from soil evaporation and transpiration of the leaves in the lower canopy (2%).

Tree species richness as an important biotic factor regulates the soil phosphorus density in China's mature natural forests.

Tree species richness has been recognized as an underlying driving factor for regulating soil phosphorus (P) status in many site-specific studies. However, it remains poorly understood whether this is true at broad scales where soil P strongly rely on climate, soil type and vegetation type. Here, based on the data of 946 mature natural forest sites from a nationwide field survey in China, we analyzed the impact of tree species richness on soil P density of China's mature natural forests (deciduous coniferous forest, DCF; evergreen coniferous forest, ECF; deciduous broad-leaved forest, DBF; evergreen broad-leaved forest, EBF; and mixed coniferous and broad-leaved forest, MF). Our results showed that tree species richness had a negative effect on soil P density in China's mature natural forests. The Random Forest regression model showed that the relative importance of tree species richness to soil P density was second only to the climate factors (mean annual temperature, MAT; mean annual precipitation, MAP). In addition, the structural equation model (SEM) results showed that the goodness fit of SEM increased when the tree species richness was included into the model. These results suggested that tree species richness was an important factor in regulating the China's mature natural forests soil P density. Furthermore, the SEM results showed that the decreased soil P density was related to the increase in ANPP and the decrease in litter P concentration induced by tree species richness. This result indicates that tree species richness could facilitate plant P absorption and inhibit plant P return into the soil, and thus reducing the soil P density in China's mature natural forests. In conclusion, we found tree species richness was an important biotic factor in regulating soil P density at broad scales, which should be fully considered in Earth models that represent P cycle.

Spatial variation and mechanisms of leaf water content in grassland plants at the biome scale: evidence from three comparative transects.

Leaf water content (LWC) has important physiological and ecological significance for plant growth. However, it is still unclear how LWC varies over large spatial scale and with plant adaptation strategies. Here, we measured the LWC of 1365 grassland plants, along three comparative precipitation transects from meadow to desert on the Mongolia Plateau (MP), Loess Plateau, and Tibetan Plateau, respectively, to explore its spatial variation and the underlying mechanisms that determine this variation. The LWC data were normally distributed with an average value of 0.66 g g-1. LWC was not significantly different among the three plateaus, but it differed significantly among different plant life forms. Spatially, LWC in the three plateaus all decreased and then increased from meadow to desert grassland along a precipitation gradient. Unexpectedly, climate and genetic evolution only explained a small proportion of the spatial variation of LWC in all plateaus, and LWC was only weakly correlated with precipitation in the water-limited MP. Overall, the lasso variation in LWC with precipitation in all plateaus represented an underlying trade-off between structural investment and water income in plants, for better survival in various environments. In brief, plants should invest less to thrive in a humid environment (meadow), increase more investment to keep a relatively stable LWC in a drying environment, and have high investment to hold higher LWC in a dry environment (desert). Combined, these results indicate that LWC should be an important variable in future studies of large-scale trait variations.