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  • Volume 19 Issue 2
    The cover image features the vast Helan Grassland, located west of the Helan Mountains in China. This arid landscape represents both a research site and a microcosm of ecological restoration across China's drylands. In recent years, local ecological initiatives, such as grazing withdrawal, the Three-North Shelterbelt Program, desertification control and ecosystem restoration, have reversed environmental degradation and curbed desert encroachment from the Tengger and Ulan Buh Deserts. In this issue, Gao et al. evaluated the effects of climate change and human activities on soil multifunctionality (SMF) in China's drylands. Their findings show that positive interventions, including fencing and the Grain for Green Program, enhance soil nutrient supply, productivity and climate regulation. Photo taken by Siyuan Gao. See Gao et al. in this issue for details
      
    Research Article
    Jiatao Zhang, Lan Du, Yonghong Luo, Yann Hautier, Ru Tian, Yan Shen, Mohsin Mahmood, Zhuwen Xu
    2026, 19 (2): rtaf167.
    Abstract ( 111 )   PDF(pc) (1516KB) ( 44 )   Save
    Drought events and nitrogen deposition are substantially modifying the stability of terrestrial ecosystems. Previous studies have mostly investigated these factors separately, with an emphasis on productivity stability, leaving their combined effects on multiple dimensions of ecosystem stability poorly understood. We conducted a four-year grassland manipulative experiment to examine how three drought scenarios—intense drought, chronic drought, and precipitation frequency reduction—interact with nitrogen addition to influence community compositional stability and productivity stability. The results showed that drought and nitrogen enrichment independently influenced grassland stability without significant interactions. Both intense and chronic drought reduced productivity stability, while reduced precipitation frequency decreased compositional stability. Nitrogen addition decreased both types of stability. Productivity stability was driven by the dominant species’ productivity stability or a combination of it and species asynchrony, depending on the drought scenario. Compositional stability consistently depended on the dominant species’ compositional stability. Compositional and productivity stability remained decoupled across treatments. This study provides the first empirical evidence of the divergent responses of grassland compositional and productivity stability to various drought scenarios under nitrogen enrichment. Our findings highlight the importance of prioritizing dominant species and promoting species coexistence with diverse environmental responses to maintain stable grassland composition and productivity under global change.
    Based on a four-year manipulative experiment, this study compared the effects of different drought scenarios and nitrogen enrichment on compositional and productivity stability in a temperate grassland. We found that these two dimensions of stability respond differently to drought types and nitrogen addition and remain persistently decoupled, highlighting the need to prioritize dominant species and promote the coexistence of species with diverse environmental response strategies to maintain grassland stability under global change.
    Deyun Chen, Zhiyun Lu, Yanping Wang, Chunyu Hou, Jianping Wu
    2026, 19 (2): rtaf175.
    Abstract ( 61 )   PDF(pc) (1606KB) ( 7 )   Save
    CO2 released into the atmosphere through soil respiration represents the second-largest carbon flux between terrestrial ecosystems and the atmosphere. While extensive research has concentrated on surface soils, limited studies have explored CO2 emission patterns and their primary drivers across varying soil depths. In this study, soil CO2 emissions were measured using static chambers at six different depths (10, 50, 100, 200, 300 and 400 cm) in a primary forest. Additionally, potential influencing factors, including soil physical and chemical properties, microbial diversity and community structure and function, were assessed. The results demonstrated that soil nutrients, along with fungal and bacterial diversity, generally declined with increasing soil depth. Soil CO2 emissions also decreased significantly with depth, driven primarily by biotic factors such as fungal and bacterial alpha diversity and abiotic factors such as ammonium nitrogen and available phosphorus. These findings provide new insights into the mechanisms of carbon cycling within deep soil layers in forest ecosystems.
    The patterns and drivers of carbon cycling within deep soil layers are important in forest ecosystems. Here the authors show the decreased soil CO2 emission along depth is driven by declining microbial diversity and nutrient availability.
    Peng Wang , Lei Li , Qiuyan Wang , Yanju Gao , Bo Zhang , Xiangyi Li , Qiang Zeng
    2026, 19 (2): rtaf210.
    Abstract ( 78 )   PDF(pc) (1896KB) ( 11 )   Save
    In water-limited ecosystems, photodegradation is a dominant pathway of carbon (C) turnover. However, the combined effects of spectral composition and litter traits in hyper-arid deserts remain poorly constrained, limiting the accuracy of C flux predictions. In a 637-day field experiment with three representative desert species (Populus euphratica, Alhagi sparsifolia and Karelinia caspia), we applied natural light filters to establish six spectral treatments. Full-spectrum exposure increased litter decomposition by 70%; of the five individual wave-bands, only UV-B (280–315 nm) and blue light (400–500 nm) significantly accelerated mass loss, accounting for 43% and 29% of the full-spectrum effect, respectively. These two wave-bands accelerated C and cellulose loss, whereas green light (500–580 nm) selectively promoted hemicellulose and lignin degradation without affecting total mass. UV-B and blue light also increased specific leaf area (SLA) by 12.9% and 7.0% and elevated litter microbial respiration rate (LMR, 24-h incubation at 25 °C, 60% relative humidity) by 54.6% and 40.6%, respectively. Across four spectral regions, initial C content, lignin:N ratio, SLA and LMR were significantly correlated with photodegradation rate, with LMR and lignin content per unit surface area being the strongest predictors of susceptibility. These results highlight the co-regulation of photodegradation by spectral composition and plant traits, advancing mechanistic understanding of C cycling in hyper-arid ecosystems and providing refined parameters for Earth system models changing solar regimes.
    This study quantified litter photodegradation for three representative Taklimakan plant species under six spectral filters. Results showed that UV-B and blue light are the primary spectral drivers of litter photodegradation, with plant traits (lignin, SLA) jointly governing the carbon-loss rate.
    Review
    Hui Jiang, Yuan-Yuan Zeng, Xu-Dong Liu, Francisco J Corpas, Md Mahadi Hasan, Feng-Ping Li, Xue-Qian Tian, Yu-Hang Zhang, Rui-Yun He, Xiang-Wen Fang
    2026, 19 (2): rtaf172.
    Abstract ( 159 )   PDF(pc) (1400KB) ( 19 )   Save
    Drought is one of the major abiotic stresses that limits plant growth. Roots, the primary organs responsible for water uptake, are the first to perceive and respond to water deficit conditions. Therefore, maintaining activity and promoting primary root elongation under drought stress are considered critical adaptive strategies for drought survival. However, despite increasing evidence of primary root elongation under drought stress, the underlying mechanisms, particularly hormonal regulation and its integration with environmental cues, remain poorly understood across different species and developmental contexts. Previous studies have shown that a highly coordinated network of plant hormone crosstalk plays a significant role in this adaptive process. In this review, data from diverse plant species confirmed the promoting effect of drought on primary root growth. This study further elucidated the role of abscisic acid (ABA) in enhancing primary root growth under drought conditions and explored the potential coordination of ABA with other hormones, including ethylene, auxin, and cytokinin, to synergistically promote primary root elongation. Furthermore, several physiological processes, such as cell cycle regulation, osmotic adjustment, and lateral root growth dynamics, had been systematically investigated for their contributions to this adaptive response. By dissecting these mechanisms, this study aims to propose feasible strategies to increase plant drought tolerance through targeted root system architecture modification, addressing challenges posed by future climate scenarios.
    Plant roots undergo elongation growth to acquire water, which is one of the crucial strategies for their adaptation to drought stress. However, the growth patterns and underlying mechanisms remain unclear. This review compiles the data from 197 plant species, confirms that moderate drought has a promoting effect on primary root growth, clarifies the critical role of abscisic acid in promoting primary root elongation, and discusses the synergistic interactions between abscisic acid, ethylene, auxin and cytokinin. This review provides an in-depth understanding of how plants improve their drought resistance by regulating root growth.
    Research Article
    Zhijie Li, Ling Xiong, Lars Vesterdal, Qiqian Wu, Josep Peñuelas, Kai Yue, Shengmin Zhang, Zimin Li, Kun Guo, Petr Heděnec, Fuzhong Wu, Yan Peng
    2026, 19 (2): rtaf147.
    Abstract ( 200 )   PDF(pc) (1942KB) ( 37 )   Save
    Tree planting is widely recognized as an effective strategy for enhancing terrestrial carbon sequestration, playing a crucial role in mitigating global climate change. However, our understanding of how it may affect soil fauna communities remains scarce. Here, we performed a global meta-analysis with 14 281 paired observations to evaluate tree planting effects on soil fauna abundance, biomass, and diversity across multiple former ecosystem types. Results showed that (i) tree planting had limited overall effects on soil fauna communities, only increasing Acari abundance, Protozoa abundance and Arthropod biomass by 36.9%, 56.9% and 777.3%, respectively, and decreasing the taxonomic richness of Collembola, the Pielou index of earthworm, and the Simpson index of Protozoa by 17.9%, 38.7%, and 77.1%, respectively; (ii) afforestation in non-forest lands showed strong positive effects on soil fauna abundance and diversity, especially in deserts where the abundance and Shannon-Wiener index of total soil fauna were increased by 92.5% and 65.8%, respectively, while reforestation in former forest lands generally had negative impacts; and (iii) tree planting effects on soil fauna were mediated by stand characteristics (e.g. stand age, canopy density, tree diameter) and pre-planting soil properties (e.g. bulk density, pH, carbon, nitrogen), but not by tree species type (leaf type or mycorrhizal association). These results demonstrate the contrasting effects of tree planting on soil fauna communities among different former ecosystem types, highlighting the importance of considering the legacy of former ecosystems when designing tree planting policies to restore/enhance carbon sequestration and biodiversity conservation under global environmental change scenarios.
    Based on 14,281 paired observations worldwide, this study quantifies the effects of tree planting on soil fauna abundance, biomass, and diversity. Results show that these effects strongly depend on former ecosystem types, with generally positive responses in non-forest ecosystems (especially deserts) but negative responses in previously forested lands.
    Yimin Zhao, Zhen Zhang, Li Zhang, Miaojun Ma, Guorui Hu, Shurong Zhou
    2026, 19 (2): rtaf158.
    Abstract ( 130 )   PDF(pc) (1412KB) ( 29 )   Save
    Extreme climatic events often co-occur with persistent environmental disturbances, such as nitrogen enrichment, which may influence the resistance and recovery of plant communities to extreme climatic conditions. However, most studies have focused on the resistance and recovery of community functions (e.g. biomass) to climatic events while neglecting the corresponding responses of diversity. Here, we performed a soil nitrogen addition experiment in an alpine meadow from 2011 to 2020, with 2015 characterized by extreme drought. We explored the effects of nitrogen addition on the resistance and recovery of plant community biomass and diversity in response to extreme drought using measures including community biomass, taxonomic diversity (TD), phylogenetic diversity (PD) and functional diversity (FD). We found that nitrogen addition decreased biomass resistance, mainly due to species asynchrony rather than the diversity resistance, even though PD and FD resistance also declined. Meanwhile, nitrogen addition enhanced the recovery of biomass to drought. This was mainly attributable to the direct, positive impact of nitrogen on biomass recovery, coupled with an indirect influence of species asynchrony, without any diversity (TD, PD, FD) recovery effects. Our results indicate that soil nitrogen enrichment mainly influences plant biomass responses to extreme drought, with a relatively small effect on plant diversity. Additionally, the mechanisms driving diversity and biomass responses may operate independently, as changes in diversity response did not scale up to changes in biomass. We anticipate that maintenance of plant community biomass during extreme drought would be more challenging in conditions of high nitrogen deposition.
    Nitrogen enrichment is known to modify ecosystem functional responses to drought, yet its effects on biodiversity responses and their coupling with ecosystem functioning remain unresolved. We show that nitrogen addition markedly altered the resistance and recovery of plant biomass to drought, while these responses were decoupled from changes in biodiversity resistance and recovery, suggesting that ecosystem functioning and biodiversity respond to drought largely independently across different dimensions.
    Research Article
    Yuyao Nan, Chuanshan Zou, Xingchang Wang, Qi Wang
    2026, 19 (2): rtaf164.
    Abstract ( 65 )   PDF(pc) (1931KB) ( 4 )   Save
    Herbivorous insects shape plant growth and community assembly, while conversely, plant traits, especially leaf traits, profoundly affect herbivore behaviors. However, which leaf traits and how they dominantly drive insect herbivory in a natural forest habitat remain undefined. In this study, we evaluated the background insect herbivory of 45 individual trees from five broadleaved tree species in the northeast of China. Based on that, 17 leaf traits representing four groups—leaf structural traits, photosynthetic pigments, nutrient traits, and secondary metabolites—were measured. Finally, the correlation between 17 leaf traits and background insect herbivory was investigated. The results indicated that the damaged leaf area (DLA) exhibited positive correlations with leaf structural traits (leaf mass per area and leaf size) and nutrient traits (soluble sugar (SS), non-structural carbohydrates (NSC), SS/NSC), and a negative correlation with photosynthetic pigments, nitrogen/phosphorus ratio, and anthocyanins. Meanwhile, SS/NSC, SS, leaf size, total phenol, and phosphorus were identified as the five relatively important leaf traits contributing to DLA by the quantitative estimation based on SHAP (SHapley Additive exPlanations) values. Furthermore, nutrient traits accounted for 52.9% of DLA explanation, showing the most important group of leaf traits. In addition, structural traits and secondary metabolites were found to interactively influence herbivory. These findings provide valuable insights into the complex interactions between host plants and herbivorous insects in forest ecosystems.
    Which leaf traits and how they dominantly drive insect herbivory in a natural forest habitat remain undefined. Investigating the relationship between 17 leaf traits representing four groups and damaged leaf area in five broadleaved tree species in Northeast China, this study finds that leaf nutrient content may play a critical role in driving herbivorous insect feeding preference.
    Mengjie Liu, Quan-Cheng Wang, Yang Li, Ronglei Zhou, Junxiao Pan, Dashuan Tian, Ruiyang Zhang, Houkun Chu, Ning Liu, Hui Wang, Huichen Zhang, Jingjing Shi, Ruifa Wang, Lei Ma, Shuli Niu, Jinsong Wang
    2026, 19 (2): rtaf146.
    Abstract ( 227 )   PDF(pc) (1555KB) ( 42 )   Save
    Peatlands store approximately one-third of global soil organic carbon (SOC) and clarifying SOC sources is essential to assess soil carbon (C) formation and stability in these C-rich ecosystems. However, large-scale patterns and drivers of plant- and microbial-derived C in peatlands remain poorly understood. This study applied lignin phenols and amino sugars as biomarkers for plant and microbial residues to investigate the regional distributions and controlling factors of plant- and microbial-derived C in surface peat (0–20 cm) across Zoige alpine peatlands. Our results showed that amino sugars contributed less while lignin phenols remained stable with SOC accrual, indicating the key role of plant-derived C in SOC accumulation. Soil nutrients and microbial properties explained the majority of the variation in lignin phenols, while soil nutrients and mineral protection played a more important role in amino sugars than microbial variables and climatic factors. Specifically, lignin phenols were negatively correlated with soil nutrients, fungal richness, and acid phosphatase activity, while showing a positive association with leucine aminopeptidase activity. In contrast, amino sugars were positively related to soil total phosphorus but negatively linked with Fe-associated C and Fe/Al-oxide. These findings provide the first empirical evidence of plant- and microbial-derived C and their divergent drivers in alpine peatlands over broad geographic scale, which advances our understanding of soil C formation and stability in these C-rich, climate-sensitive ecosystems.
    This study systematically investigated large-scale patterns and drivers of plant- and microbial-derived carbon (C) in soils of the Zoige alpine peatlands. It revealed that the spatial variation of lignin phenols (plant-derived C) was primarily influenced by soil nutrients and microbial properties, while amino sugars (microbial-derived C) were mainly affected by soil nutrients and mineral protection. These findings provide the first empirical evidence of plant- and microbial-derived C and their divergent drivers in alpine peatlands over a broad scale, which advances our understanding of soil C formation and stability in these C-rich, climate-sensitive ecosystems.
    Rong Hu, Mengxuan Wang, Aixin Hou, Xiaohong Guo, Xiang Long, Ying Xu, Feilong Huang, Sifan Peng, Fanzhu Qu, Xuehong Wang, Zhikang Wang, Yating Li, Bo Guan
    2026, 19 (2): rtaf169.
    Abstract ( 84 )   PDF(pc) (2295KB) ( 13 )   Save
    Arbuscular mycorrhizal fungi (AMF) can enhance terrestrial plant growth by promoting nutrient uptake. However, the effects of interactions between AMF and nutrient inputs on plant functional traits and their trade-offs remain poorly understood. In this study, a pot experiment was conducted using Phragmites australis as the host species, with AMF inoculation and non-inoculation treatments under three levels of nitrogen addition and two levels of phosphorus addition. The results showed that AMF inoculation significantly increased AMF colonization in the roots of P. australis. Under unfertilized conditions, AMF significantly promoted plant morphological development and biomass accumulation. Nitrogen addition primarily enhanced aboveground growth, while phosphorus addition significantly stimulated root system development. Regarding photosynthetic traits, AMF inoculation significantly increased carotenoid content, whereas phosphorus addition significantly reduced net photosynthetic rate (Pn), transpiration rate (Tr), and stomatal conductance (Gs) by 18.5%, 25.8%, and 28.2%, respectively. Furthermore, AMF inoculation increased leaf nitrogen and phosphorus contents but reduced stem nitrogen content and the N:P ratios in fine roots, rhizomes, and stems. Trait network analysis revealed that AMF inoculation shifted the central traits from leaf biomass to rhizome biomass, indicating a transition in hub traits from aboveground to belowground organs. These findings suggest that AMF inoculation significantly promotes the growth of P. australis although the magnitude of this benefit is modulated by soil nitrogen and phosphorus availability, with stronger effects observed under nutrient-poor conditions. This study advances our understanding of nutrient–AMF–plant interactions in wetland ecosystems and provides a theoretical foundation for ecological restoration and nutrient management in vulnerable aquatic habitats.
    This study highlights how Arbuscular mycorrhizal fungi (AMF) inoculation influences the functional traits of Phragmites australis under different nutrient conditions. It demonstrates that AMF inoculation significantly enhance growth and nutrient acquisition under low nutrients, but their benefits weaken with fertilization, shifting the trait network focus from aboveground to belowground organs.
    Na Li, Suke Hou, Mai-He Li, Xi Qi, Liangjun Zhu, Jitang Li, Paolo Cherubini, Zhaoyong Hu, Zhengfang Wu, Renkai Dong, Lei Wang, Youwen Wang, Danyang Yuan, Haibo Du
    2026, 19 (2): rtaf181.
    Abstract ( 65 )   PDF(pc) (2162KB) ( 8 )   Save
    As global warming drives plant upward migration, the alpine tundra of Changbai Mountain is experiencing encroachment by Deyeuxia angustifolia (Komarov) Y. L. Chang, a low-elevation herb. However, its impact on native shrubs such as Rhododendron aureum Georgi remains unclear. Here, we analyzed the radial growth trends and climate sensitivity of R. aureum across elevations and encroachment gradients using linear and mixed-effects model methods, and explored the mediating roles of soil properties and plant traits. Our study revealed that R. aureum exhibited stronger positive long-term growth trend at higher elevations compared to lower elevations. Mild and moderate encroachment of D. angustifolia enhanced the positive growth trend of R. aureum, especially at the low elevations. Moreover, R. aureum showed weak climate sensitivity at mid-elevation but stronger responses to winter temperatures at low elevation and to spring–summer temperatures and precipitation at high elevation. D. angustifolia encroachment further intensified this sensitivity, characterized by stronger negative responses to spring, autumn, and winter temperatures but positive responses to summer temperatures and autumn precipitation. Overall, elevation primarily influenced R. aureum growth and its sensitivity to precipitation through soil conditions and plant size traits. Soil conditions and leaf economic traits influence temperature sensitivity. These findings advance understanding of alpine vegetation dynamics and contribute to ecosystem conservation under climate change.
    Increasing elevation and herbaceous encroachment synergistically enhanced the growth trend and climate sensitivity of the alpine tundra shrub Rhododendron aureum on Changbai Mountain, with a stronger encroachment effect observed at lower elevations. Rising elevation and herbaceous encroachment jointly enhanced the growth trend and climate sensitivity of the alpine tundra shrub Rhododendron aureum on Changbai Mountain, with the encroachment effect being more pronounced at lower elevations. The findings deepen our understanding of alpine vegetation shifts and invasion mechanisms under climate change, offering scientific guidance for ecosystem conservation and management in alpine tundra regions.
    Tiancai Zhou, Gusang Qunzong, Jian Sun
    2026, 19 (2): rtaf176.
    Abstract ( 60 )   PDF(pc) (1542KB) ( 28 )   Save
    Despite nitrogen (N) and phosphorus (P) being biologically coupled and controlling many biochemical reactions, few studies have examined the N:P patterns and controls of legumes and non-legumes at a global scale. Herein, we explored how the ratio of N and P in legumes and non-legumes responds to environmental factors, globally. Our results indicated that legumes exhibited stronger N-P coupling (R² = 0.39, P < 0.0001) in warm-humid environments (mean precipitation: 988.94 mm, temperature: 12.63 °C), and the N:P is negatively affected by the soil total P (scored at = −0.25). In contrast, non-legumes were more flexible in N and P (R2 = 0.23, P < 0.0001) in semihumid regions (precipitation = 785.01 mm, temperature = 8.85 °C), where soil total N (scored at = −0.22) and biodiversity (scored at = 0.16) emerge as dominant drivers for the N:P. Although legumes are expected to be more soil P-limited, our findings revealed that the leaf N and P were more coupled in legumes than in non-legumes, which offers a unique perspective on resource utilization and survival strategies in different plant functions.
    Although legumes are expected to be more soil P-limited, our findings reveal that the leaf N and P are more coupled in legumes than in non-legumes, which offers a unique perspective on resource utilization and survival strategies in different plant functions.
    Short Communication
    Senhao Wang, Kaijun Zhou, Taiki Mori, Andi Li, Cheng Peng, Guangcan Yu, Enqing Hou, Sihan Yao, Linjie Fan, Baixin Zhang, Yufang Wang, Qing Ye, Weiren Wang, Shun Zou, Xiangping Tan, Qinggong Mao, Mianhai Zheng, Juan Huang, Zhanfeng Liu, Jiangming Mo, Wei Zhang
    2026, 19 (2): rtaf163.
    Abstract ( 109 )   PDF(pc) (750KB) ( 16 )   Save
    Leguminous species have an advantage in acquiring phosphorus (P) compared with non-leguminous species. Nonetheless, it remains unclear whether this advantage would diminish under long-term nitrogen (N) deposition. In the seventh year of a simulated long-term N deposition experiment, we sampled surface soil to measure acid phosphatase activity (ACP) in stands dominated by leguminous and non-leguminous species, respectively. To assess the response of ACP to prolonged N addition, we also collected data on ACP in the second, eighth, twelfth, and thirteenth years of the experiment. We found that the difference in soil ACP between the two plantation types disappeared after long-term N input, and this process was accelerated under high N addition. This occurred due to an exacerbated P-limitation, which primarily prompts ACP production only in non-leguminous species. Additionally, there was a relatively decreased N contribution efficiency to ACP in the stand with leguminous species, indicating that soil N content no longer primarily governs ACP. This study demonstrates that prolonged high N deposition accelerates the loss of the “P-acquiring advantage” in leguminous plantations. To elucidate the differences in P usage strategies between leguminous and non-leguminous species under global change, more systematic research is warranted in the future.
    Whether the “phosphorus-acquiring advantage” of legumes in subtropical regions can persist under long-term nitrogen deposition remains unclear. Here, we demonstrate that prolonged high-nitrogen inputs accelerate the loss of this advantage in legume plantations.
    Correction
IF: 3.9
CiteScore: 5.7
Editors-in-Chief
Yuanhe Yang
Bernhard Schmid
CN 10-1172/Q
ISSN 1752-9921(print)
ISSN 1752-993X(online)