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.

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.

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.

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.

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 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.

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.

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.

Mapping forest type and age in China's plantations.

Forest age serves as an essential factor in determining the accuracy of historical and future carbon (C) uptake quantifications, which is especially critical for China since the forest C stock dynamics are sensitive to the fast-growing, young-age plantations. However, a spatially explicit forest age maps with specific focus on forest plantations is not available yet. In this study, we developed a 1-km resolution age and type maps of forest plantations, and quantified their uncertainties spatially using field-measured data, national forest inventory data, digitalized forest maps, and remote sensing-based forest height maps. Simulation results showed forest plantations were 16.5 years old at national scale in 2005, which is close to the age of 16.6 years old derived from the 7th national inventory data using medium age in each forest plantation group with weighted area. Interestingly, we found that human management played an important role in forest age map reconstruction, which has not yet been considered in former studies. We also suggest that forest age and type maps should be used consistently in C stock simulations to avoid biases from mismatch information. Large uncertainty found in this study suggests further endeavors are required for improving the forest age and type maps.

Plant Trait Networks: Improved Resolution of the Dimensionality of Adaptation.

Functional traits are frequently used to evaluate plant adaptation across environments. Yet, traits tend to have multiple functions and interactions, which cannot be accounted for in traditional correlation analyses. Plant trait networks (PTNs) clarify complex relationships among traits, enable the calculation of metrics for the topology of trait coordination and the importance of given traits in PTNs, and how they shift across communities. Recent studies of PTNs provide new insights into some important topics, including trait dimensionality, trait spectra (including the leaf economic spectrum), stoichiometric principles, and the variation of phenotypic integration along gradients of resource availability. PTNs provide improved resolution of the multiple dimensions of plant adaptation across scales and responses to shifting resources, disturbance regimes, and global change.

Diversity-decomposition relationships in forests worldwide.

Plant species diversity affects carbon and nutrient cycling during litter decomposition, yet the generality of the direction of this effect and its magnitude remains uncertain. With a meta-analysis including 65 field studies across the Earth's major forest ecosystems, we show here that decomposition was faster when litter was composed of more than one species. These positive biodiversity effects were mostly driven by temperate forests but were more variable in other forests. Litter mixture effects emerged most strongly in early decomposition stages and were related to divergence in litter quality. Litter diversity also accelerated nitrogen, but not phosphorus release, potentially indicating a decoupling of nitrogen and phosphorus cycling and perhaps a shift in ecosystem nutrient limitation with changing biodiversity. Our findings demonstrate the importance of litter diversity effects for carbon and nutrient dynamics during decomposition, and show how these effects vary with litter traits, decomposer complexity and forest characteristics.

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.

Climate and litter C/N ratio constrain soil organic carbon accumulation.

Soil organic carbon (SOC) plays critical roles in stabilizing atmospheric CO2 concentration, but the mechanistic controls on the amount and distribution of SOC on global scales are not well understood. In turn, this has hampered the ability to model global C budgets and to find measures to mitigate climate change. Here, based on the data from a large field survey campaign with 2600 plots across China's forest ecosystems and a global collection of published data from forested land, we find that a low litter carbon-to-nitrogen ratio (C/N) and high wetness index (P/PET, precipitation-to-potential-evapotranspiration ratio) are the two factors that promote SOC accumulation, with only minor contributions of litter quantity and soil texture. The field survey data demonstrated that high plant diversity decreased litter C/N and thus indirectly promoted SOC accumulation by increasing the litter quality. We conclude that any changes in plant-community composition, plant-species richness and environmental factors that can reduce the litter C/N ratio, or climatic changes that increase wetness index, may promote SOC accumulation. The study provides a guideline for modeling the carbon cycle of various ecosystem scales and formulates the principle for land-based actions for mitigating the rising atmospheric CO2 concentration.

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).

Effects of sky conditions on net ecosystem productivity of a subtropical coniferous plantation vary from half-hourly to daily timescales.

The dynamic changes of solar radiation have received wide attention in global change studies, but there are controversies about the influence of diffuse radiation on ecosystem carbon sequestration. Using eddy covariance measurements from 2010 to 2012, the effects of sky conditions extracted from adjacent sunny, cloudy, and overcast days on net ecosystem productivity (NEP) of a subtropical coniferous plantation were examined from half-hourly to daily scales. Half-hourly NEP responded to the changing radiation more efficiently on overcast days compared to sunny days, but such response did not differ obviously between cloudy and sunny days. Compared with sunny conditions, apparent quantum yield (α) under overcast (cloudy) conditions changed 282.4% (41.7%) in spring, 140.3% (-4.2%) in summer, 218.5% (38.9%) in autumn, and 146.2% (0.5%) in winter, respectively; annually, α under overcast (cloudy) conditions increased by 225.9% (19.8%) in 2010, 189.8% (6.0%) in 2011, and 159.5% (21.4%) in 2012, respectively. Moreover, the potential NEP at the light intensity of 150 and 750 W m-2 was improved due to increased diffuse fraction. However, both daytime NEP and daily NEP were significantly lower under overcast skies than under sunny and cloudy skies. Compared with sunny days, daily NEP on overcast days decreased by 127.7% in spring, 126.4% in summer, 121.8% in autumn, and 100.6% in winter, respectively; annually, daily NEP decreased by 122.5% in 2010, 141.7% in 2011, and 109.9% in 2012, respectively. Diurnal patterns of daily NEP were quite similar between sunny and cloudy days. Both path analysis and multiple regression showed that solar radiation, especially diffuse radiation, was responsible for the variations of NEP under different skies across seasons, but this effect may be weakened by seasonal droughts. This study implies that the effects of sky conditions on NEP are timescale dependent and should be paid more attention in ecosystem carbon cycle study.

Ecosystem Traits Linking Functional Traits to Macroecology.

As the range of studies on macroecology and functional traits expands, integration of traits into higher-level approaches offers new opportunities to improve clarification of larger-scale patterns and their mechanisms and predictions using models. Here, we propose a framework for quantifying 'ecosystem traits' and means to address the challenges of broadening the applicability of functional traits to macroecology. Ecosystem traits are traits or quantitative characteristics of organisms (plants, animals, and microbes) at the community level expressed as the intensity (or density) normalized per unit land area. Ecosystem traits can inter-relate and integrate data from field trait surveys, eddy-flux observation, remote sensing, and ecological models, and thereby provide new resolution of the responses and feedback at regional to global scale.