A continental-scale analysis reveals the latitudinal gradient of stomatal density across amphistomatous species: Evolutionary history vs. present-day environment.

BACKGROUND AND AIMS: Amphistomy is a potential method for increasing photosynthetic rate; however, the latitudinal gradients of stomatal density across amphistomatous species and their drivers remain unknown. METHODS: Here, the adaxial stomatal density (SDad) and abaxial stomatal density (SDab) of 486 amphistomatous species-site combinations, belonging to 32 plant families, were collected from China, and their total stomatal density (SDtotal) and stomatal ratio (SR) were calculated. KEY RESULTS: Overall, these four stomatal traits did not show significant phylogenetic signals. There were no significant differences in SDab and SDtotal between woody and herbaceous species, but SDad and SR were higher in woody species than in herbaceous species. Besides, a significantly positive relationship between SDab and SDad was observed. We also found that stomatal density (including SDab, SDad, and SDtotal) decreased with latitude while SR increased with latitude, and temperature seasonality was the most important environmental factor driving it. Besides, evolutionary history (represented by both phylogeny and species) explained about 10-22 fold more of the variation in stomatal traits than the present-day environment (65.2%-71.1% vs. 2.9%-6.8%). CONCLUSIONS: Our study extended our knowledge of trait-environment relationships and highlighted the importance of evolutionary history in driving stomatal trait variability.

Divergent response and adaptation of specific leaf area to environmental change at different spatio-temporal scales jointly improve plant survival.

Specific leaf area (SLA) is one of the most important plant functional traits. It integrates multiple functions and reflects strategies of plants to obtain resources. How plants employ different strategies (e.g., through SLA) to respond to dynamic environmental conditions remains poorly understood. This study aimed to explore the spatial variation in SLA and its divergent adaptation through the lens of biogeographic patterns, evolutionary history, and short-term responses. SLA data for 5424 plant species from 76 natural communities in China were systematically measured and integrated with meta-analysis of field experiments (i.e., global warming, drought, and nitrogen addition). The mean value of SLA across all species was 21.8 m2  kg-1 , ranging from 0.9 to 110.2 m2  kg-1 . SLA differed among different ecosystems, temperature zones, vegetation types, and functional groups. Phylogeny had a weak effect on SLA, but plant species evolved toward higher SLA. Furthermore, SLA responded nonlinearly to environmental change. Unexpectedly, radiation was one of the main factors determining the spatial variation in SLA on a large scale. Conversely, short-term manipulative experiments showed that SLA increased with increased resource availability and tended to stabilize with treatment duration. However, different species exhibited varying response patterns. Overall, variation in long-term adaptation of SLA to environmental gradients and its short-term response to resource pulses jointly improve plant adaptability to a changing environment. Overall SLA-environment relationships should be emphasized as a multidimensional strategy for elucidating environmental change in future research.

The essential role of biodiversity in the key axes of ecosystem function.

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.

The potential bias of nitrogen deposition effects on primary productivity and biodiversity.

Atmospheric nitrogen (N) deposition is composed of both inorganic nitrogen (IN) and organic nitrogen (ON), and these sources of N may exhibit different impacts on ecosystems. However, our understanding of the impacts of N deposition is largely based on experimental gradients of INs or more rarely ONs. Thus, the effects of N deposition on ecosystem productivity and biodiversity may be biased. We explored the differential impacts of N addition with different IN:ON ratios (0:10, 3:7, 5:5, 7:3, and 10:0) on aboveground net primary productivity (ANPP) of plant community and plant diversity in a typical temperate grassland with a long-term N addition experiment. Soil pH, litter biomass, soil IN concentration, and light penetration were measured to examine the potential mechanisms underlying species loss with N addition. Our results showed that N addition significantly increased plant community ANPP by 68.33%-105.50% and reduced species richness by 16.20%-37.99%. The IN:ON ratios showed no significant effects on plant community ANPP. However, IN-induced species richness loss was about 2.34 times of ON-induced richness loss. Soil pH was positively related to species richness, and they exhibited very similar response patterns to IN:ON ratios. It implies that soil acidification accounts for the different magnitudes of species loss with IN and ON additions. Overall, our study suggests that it might be reasonable to evaluate the effects of N deposition on plant community ANPP with either IN or ON addition. However, the evaluation of N deposition on biodiversity might be overestimated if only IN is added or underestimated if only ON is added.

High environmental costs behind rapid economic development: Evidence from economic loss caused by atmospheric acid deposition.

The rapid growth of energy-intensive and high-emission industries has propelled China's economy but has also led to massive levels of air pollutant emissions and ecological problems, such as acid deposition. Despite recent declines, atmospheric acid deposition in China is still severe. Long-term exposure to high levels of acid depositions has a substantial negative impact on the ecosystem. Evaluating these hazards and incorporating this issue into planning and decision-making processes is critical to achieving sustainable development goals in China. However, the long-term economic loss caused by atmospheric acid deposition and its temporal and spatial variation in China is unclear. Hence, the aim of this study was to assess the environmental cost of acid deposition in the agriculture, forestry, construction, and transportation industries from 1980 to 2019, using long-term monitoring, integrated data, and the dose-response method with localization parameters. The results showed that the estimated cumulative environmental cost of acid deposition was USD 230 billion, representing 0.27% of the gross domestic product (GDP) in China. This cost, was particularly high for building materials, followed by crops, forests, and roads. Temporally, the environmental cost and the ratio of environmental cost to GDP decreased from their peaks by 43% and 91%, respectively, because of emission controls targeting acidifying pollutants and promotion of clean energy. Spatially, the largest environmental cost occurred in developing provinces, indicating that more stringent emission reduction measures should be implemented in these regions. These findings highlight the huge environmental costs behind rapid development; however, the implementation of reasonable emission reduction measures can effectively reduce these environmental costs, providing a promising paradigm for other undeveloped and developing countries.

Leaf trait network architecture shifts with species-richness and climate across forests at continental scale.

Variation in the architecture of trait networks among ecosystems has been rarely quantified, but can provide high resolution of the contrasting adaptation of the whole phenotype. We constructed leaf trait networks (LTNs) from 35 structural, anatomical and compositional leaf traits for 394 tree species in nine forests from tropical to cold-temperate zones in China. Our analyses supported the hypothesis that LTNs would increase in modular complexity across forests in parallel with species-richness and climatic warmth and moisture, due to reduced phenotypic constraints and greater opportunities for niche differentiation. Additionally, we found that within LTNs, leaf economics traits including leaf thickness would have central importance, acting as hub traits with high connectivity due to their contributions to multiple functions. Across the continent, the greater species richness and trait diversity observed in forests under resource-rich climates enable greater complexity in whole phenotype structure and function as indicated by the trait network architecture.

The adjustment of life history strategies drives the ecological adaptations of soil microbiota to aridity.

Soil microbiota increase their fitness to local habitats by adjusting their life history strategies. Yet, how such adjustments drive their ecological adaptations in xeric grasslands remains elusive. In this study, shifts in the traits that potentially represent microbial life history strategies were studied along two aridity gradients with different climates using metagenomic and trait-based approaches. The results indicated that resource acquisition (e.g., higher activities of ß-d-glucosidase and N-acetyl-ß-d-glucosidase, higher degradation rates of cellulose and chitin, as well as genes involved in cell motility, biodegradation, transportation and competition) and growth yield (e.g., higher biomass and respiration) strategies were depleted at higher aridity. However, maintenance of cellular and high growth potential (e.g., higher metabolic quotients and genes related to DNA replication, transcription, translation, central carbon metabolism and biosynthesis) and stress tolerance (e.g., genes involved in DNA damage repair, cation transportation, sporulation and osmolyte biosynthesis) strategies were enriched at higher aridity. This implied that microbiota have lower growth yields but are probably well primed for rapid responses to pulses of rainfall in more arid soils, whereas those in less arid soils may have stronger resource acquisition and growth yield abilities. By integrating a large amount of evidence from taxonomic, metagenomic, genomic and biochemical investigations, this study demonstrates that the ecological adaptations of soil microbiota to aridity made by adjusting and optimizing their life history strategies are universal in xeric grasslands and provides an underlying mechanistic understanding of soil microbial responses to climate changes.

Contrasting adaptation and optimization of stomatal traits across communities at continental scale.

Shifts in stomatal trait distributions across contrasting environments and their linkage with ecosystem productivity at large spatial scales have been unclear. Here, we measured the maximum stomatal conductance (g), stomatal area fraction (f), and stomatal space-use efficiency (e, the ratio of g to f) of 800 plant species ranging from tropical to cold-temperate forests, and determined their values for community-weighted mean, variance, skewness, and kurtosis. We found that the community-weighted means of g and f were higher in drier sites, and thus, that drought 'avoidance' by water availability-driven growth pulses was the dominant mode of adaptation for communities at sites with low water availability. Additionally, the variance of g and f was also higher at arid sites, indicating greater functional niche differentiation, whereas that for e was lower, indicating the convergence in efficiency. When all other stomatal trait distributions were held constant, increasing kurtosis or decreasing skewness of g would improve ecosystem productivity, whereas f showed the opposite patterns, suggesting that the distributions of inter-related traits can play contrasting roles in regulating ecosystem productivity. These findings demonstrate the climatic trends of stomatal trait distributions and their significance in the prediction of ecosystem productivity.

Atmospheric Wet Iron, Molybdenum, and Vanadium Deposition in Chinese Terrestrial Ecosystems.

Iron (Fe), molybdenum (Mo), and vanadium (V) are the main components of the three known biological nitrogenases, which constrain nitrogen fixation and affect ecosystem productivity. Atmospheric deposition is an important pathway of these trace metals into ecosystems. Here, we explored the deposition flux, spatiotemporal pattern, and influencing factors of atmospheric wet Fe, Mo, and V deposition based on China Wet Deposition Observation Network (ChinaWD) data from 2016 to 2020. Our results showed that atmospheric wet Fe, Mo, and V deposition was 7.77 ± 7.24, 0.16 ± 0.11, and 0.13 ± 0.12 mg m-2 a-1 in Chinese terrestrial ecosystems, respectively, and revealed obvious spatial patterns but no significant annual trends. Wet Fe deposition was significantly correlated with the soil Fe content. Mo and V deposition was more affected by anthropogenic activities than Fe deposition. Wet Mo deposition was significantly affected by Mo ore reserves and waste incineration. V deposition was significantly correlated with domestic biomass burning. This study quantified wet Fe, Mo, and V deposition in China for the first time, and the implications of atmospheric trace metal deposition on biological nitrogen fixation were discussed.

Atmospheric silicon wet deposition and its influencing factors in China.

Silicon (Si) is considered a "quasi-essential" nutrient element for plants and is also an essential nutrient for some phytoplankton. Except for the silicate provided by weathering, atmospheric deposition has gradually become an important supplementary method for Si nutrients to enter the ecosystem. However, national observational studies on atmospheric silicon deposition have not yet been reported. Herein, based on the China Wet Deposition Observation Network, we continuously collected monthly wet deposition samples from 43 typical ecosystems from 2013 to 2020 and measured the content of dissolved silica (dSi) in precipitation to quantify the spatiotemporal patterns of Si wet deposition in China. The results showed that the mean annual dSi wet deposition in China during 2013-2020 was approximately 2.07 ± 0.27 kg ha-1 yr-1. Atmospheric dSi deposition was higher in Southwest, North, and South China but lower in the Northwest and Northeast China, which was mainly regulated by precipitation and soil available Si content. There was no significant annual variation trend in dSi deposition during 2013-2020 in China, which showed disorderly fluctuations from year to year. This study revealed the spatiotemporal patterns of atmospheric dSi deposition in China for the first time, which can provide unique scientific data to explore the potential effect of dSi deposition on carbon sequestration in aquatic ecosystems. A comprehensive evaluation of the nutrient balance of aquatic ecosystems from the perspective of nitrogen, phosphorus, and silicon stoichiometry is required in the future.

Microbial metabolic response to winter warming stabilizes soil carbon.

Current consensus on global climate change predicts warming trends with more pronounced temperature changes in winter than summer in the Northern Hemisphere at high latitudes. Moderate increases in soil temperature are generally related to faster rates of soil organic carbon (SOC) decomposition in Northern ecosystems, but there is evidence that SOC stocks have remained remarkably stable or even increased on the Tibetan Plateau under these conditions. This intriguing observation points to altered soil microbial mediation of carbon-cycling feedbacks in this region that might be related to seasonal warming. This study investigated the unexplained SOC stabilization observed on the Tibetan Plateau by quantifying microbial responses to experimental seasonal warming in a typical alpine meadow. Ecosystem respiration was reduced by 17%-38% under winter warming compared with year-round warming or no warming and coincided with decreased abundances of fungi and functional genes that control labile and stable organic carbon decomposition. Compared with year-round warming, winter warming slowed macroaggregate turnover rates by 1.6 times, increased fine intra-aggregate particulate organic matter content by 75%, and increased carbon stabilized in microaggregates within stable macroaggregates by 56%. Larger bacterial "necromass" (amino sugars) concentrations in soil under winter warming coincided with a 12% increase in carboxyl-C. These results indicate the enhanced physical preservation of SOC under winter warming and emphasize the role of soil microorganisms in aggregate life cycles. In summary, the divergent responses of SOC persistence in soils exposed to winter warming compared to year-round warming are explained by the slowing of microbial decomposition but increasing physical protection of microbially derived organic compounds. Consequently, the soil microbial response to winter warming on the Tibetan Plateau may cause negative feedbacks to global climate change and should be considered in Earth system models.

Effect of atmospheric nitrogen deposition and its components on carbon flux in terrestrial ecosystems in China.

Long-term atmospheric nitrogen (N) deposition increases bioavailable N in terrestrial ecosystems, thereby influencing ecosystem productivity. However, how N deposition and its components (i.e., NO3--N and NH4+-N) influence the spatial pattern of productivity in terrestrial ecosystems in China remains unknown. Here, we utilize published data including carbon (C) fluxes from eddy flux tower (gross ecosystem productivity, ecosystem respiration, and net ecosystem productivity) and the corresponding climate and N deposition data for 60 typical ecosystems in China. The objective was to investigate the effect of N deposition on ecosystem productivity and explore the variations of N use efficiency (NUE). Our results reveal that atmospheric total N deposition is significantly correlated with C fluxes of terrestrial ecosystems in China. Ecosystems respond variably to different components of N deposition. In detail, forest ecosystem marginally correlated with NO3--N and wet deposition, while grassland ecosystem significantly correlated with NH4+-N and dry deposition. NUE of productivity induced by N deposition in Chinese terrestrial ecosystems was 53.95 ± 40.30 g C g-1 N, and it was influenced by precipitation and aridity index. This study quantifies the contribution of total N deposition and its associated components to productivity in terrestrial ecosystems in China, offering vital information for regional C and N management.

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.

Patterns of plant carbon, nitrogen, and phosphorus concentration in relation to productivity in China's terrestrial ecosystems.

Plant nitrogen (N) and phosphorus (P) content regulate productivity and carbon (C) sequestration in terrestrial ecosystems. Estimates of the allocation of N and P content in plant tissues and the relationship between nutrient content and photosynthetic capacity are critical to predicting future ecosystem C sequestration under global change. In this study, by investigating the nutrient concentrations of plant leaves, stems, and roots across China's terrestrial biomes, we document large-scale patterns of community-level concentrations of C, N, and P. We also examine the possible correlation between nutrient content and plant production as indicated by vegetation gross primary productivity (GPP). The nationally averaged community concentrations of C, N, and P were 436.8, 14.14, and 1.11 mg·g-1 for leaves; 448.3, 3.04 and 0.31 mg·g-1 for stems; and 418.2, 4.85, and 0.47 mg·g-1 for roots, respectively. The nationally averaged leaf N and P productivity was 249.5 g C GPP·g-1 N·y-1 and 3,157.9 g C GPP·g-1 P·y-1, respectively. The N and P concentrations in stems and roots were generally more sensitive to the abiotic environment than those in leaves. There were strong power-law relationships between N (or P) content in different tissues for all biomes, which were closely coupled with vegetation GPP. These findings not only provide key parameters to develop empirical models to scale the responses of plants to global change from a single tissue to the whole community but also offer large-scale evidence of biome-dependent regulation of C sequestration by nutrients.

Effects of national ecological restoration projects on carbon sequestration in China from 2001 to 2010.

The long-term stressful utilization of forests and grasslands has led to ecosystem degradation and C loss. Since the late 1970s China has launched six key national ecological restoration projects to protect its environment and restore degraded ecosystems. Here, we conducted a large-scale field investigation and a literature survey of biomass and soil C in China's forest, shrubland, and grassland ecosystems across the regions where the six projects were implemented (∼16% of the country's land area). We investigated the changes in the C stocks of these ecosystems to evaluate the contributions of the projects to the country's C sink between 2001 and 2010. Over this decade, we estimated that the total annual C sink in the project region was 132 Tg C per y (1 Tg = 1012 g), over half of which (74 Tg C per y, 56%) was attributed to the implementation of the projects. Our results demonstrate that these restoration projects have substantially contributed to CO2 mitigation in China.

Variation and evolution of C:N ratio among different organs enable plants to adapt to N-limited environments.

Carbon (C) and nitrogen (N) are the primary elements involved in the growth and development of plants. The C:N ratio is an indicator of nitrogen use efficiency (NUE) and an input parameter for some ecological and ecosystem models. However, knowledge remains limited about the convergent or divergent variation in the C:N ratios among different plant organs (e.g., leaf, branch, trunk, and root) and how evolution and environment affect the coefficient shifts. Using systematic measurements of the leaf-branch-trunk-root of 2,139 species from tropical to cold-temperate forests, we comprehensively evaluated variation in C:N ratio in different organs in different taxa and forest types. The ratios showed convergence in the direction of change but divergence in the rate of change. Plants evolved toward lower C:N ratios in the leaf and branch, with N playing a more important role than C. The C:N ratio of plant organs (except for the leaf) was constrained by phylogeny, but not strongly. Both the change of C:N during evolution and its spatial variation (lower C:N ratio at midlatitudes) help develop the adaptive growth hypothesis. That is, plants with a higher C:N ratio promote NUE under strong N-limited conditions to ensure survival priority, whereas plants with a lower C:N ratio under less N-limited environments benefit growth priority. In nature, larger proportion of species with a high C:N ratio enabled communities to inhabit more N-limited conditions. Our results provide new insights on the evolution and drivers of C:N ratio among different plant organs, as well as provide a quantitative basis to optimize land surface process models.

Nitrogen addition does not reduce the role of spatial asynchrony in stabilising grassland communities.

While nitrogen (N) amendment is known to affect the stability of ecological communities, whether this effect is scale-dependent remains an open question. By conducting a field experiment in a temperate grassland, we found that both plant richness and temporal stability of community biomass increased with spatial scale, but N enrichment reduced richness and stability at the two scales considered. Reduced local-scale stability under N enrichment arose from N-induced reduction in population stability, which was partly attributable to the decline in local species richness, as well as reduction in asynchronous local population dynamics across species. Importantly, N enrichment did not alter spatial asynchrony among local communities, which provided similar spatial insurance effects at the larger scale, regardless of N enrichment levels. These results suggest that spatial variability among local communities, in addition to local diversity, may help stabilise ecosystems at larger spatial scales even in the face of anthropogenic environmental changes.

Microbes drive global soil nitrogen mineralization and availability.

Soil net nitrogen mineralization rate (Nmin ), which is critical for soil nitrogen availability and plant growth, is thought to be primarily controlled by climate and soil physical and/or chemical properties. However, the role of microbes on regulating soil Nmin has not been evaluated on the global scale. By compiling 1565 observational data points of potential net Nmin from 198 published studies across terrestrial ecosystems, we found that Nmin significantly increased with soil microbial biomass, total nitrogen, and mean annual precipitation, but decreased with soil pH. The variation of Nmin was ascribed predominantly to soil microbial biomass on global and biome scales. Mean annual precipitation, soil pH, and total soil nitrogen significantly influenced Nmin through soil microbes. The structural equation models (SEM) showed that soil substrates were the main factors controlling Nmin when microbial biomass was excluded. Microbe became the primary driver when it was included in SEM analysis. SEM with soil microbial biomass improved the Nmin prediction by 19% in comparison with that devoid of soil microbial biomass. The changes in Nmin contributed the most to global soil NH4+ -N variations in contrast to climate and soil properties. This study reveals the complex interactions of climate, soil properties, and microbes on Nmin and highlights the importance of soil microbial biomass in determining Nmin and nitrogen availability across the globe. The findings necessitate accurate representation of microbes in Earth system models to better predict nitrogen cycle under global change.

Joint structural and physiological control on the interannual variation in productivity in a temperate grassland: A data-model comparison.

Given the important contributions of semiarid region to global land carbon cycle, accurate modeling of the interannual variability (IAV) of terrestrial gross primary productivity (GPP) is important but remains challenging. By decomposing GPP into leaf area index (LAI) and photosynthesis per leaf area (i.e., GPP_leaf), we investigated the IAV of GPP and the mechanisms responsible in a temperate grassland of northwestern China. We further assessed six ecosystem models for their capabilities in reproducing the observed IAV of GPP in a temperate grassland from 2004 to 2011 in China. We observed that the responses to LAI and GPP_leaf to soil water significantly contributed to IAV of GPP at the grassland ecosystem. Two of six models with prescribed LAI simulated of the observed IAV of GPP quite well, but still underestimated the variance of GPP_leaf, therefore the variance of GPP. In comparison, simulated pattern by the other four models with prognostic LAI differed significantly from the observed IAV of GPP. Only some models with prognostic LAI can capture the observed sharp decline of GPP in drought years. Further analysis indicated that accurately representing the responses of GPP_leaf and leaf stomatal conductance to soil moisture are critical for the models to reproduce the observed IAV of GPP_leaf. Our framework also identified that the contributions of LAI and GPP_leaf to the observed IAV of GPP were relatively independent. We conclude that our framework of decomposing GPP into LAI and GPP_leaf has a significant potential for facilitating future model intercomparison, benchmarking and optimization should be adopted for future data-model comparisons.