Asymmetric responses of roots and hyphae to inorganic and organic nutrient additions: a global meta-analysis

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Nevertheless, the mechanisms by which roots and hyphae respond to variations in inorganic and organic nutrients remain inadequately understood. Methods We conducted a meta-analysis on the effects of nitrogen, phosphorus, nitrogen-phosphorus combination and organic nutrient additions on root length (RL), root biomass (RB), root colonization (RC), hyphal length (HL), and hyphal biomass (HB) based on 738 root-hyphae observation pairs. Results We found that RB and RL increased by 27%-34% and 31%-36%, respectively, following nitrogen and/or phosphorus additions, whereas HL increased by 26% and RC decreased by 20%. The effect sizes of organic nutrient addition on hyphae were two to three times greater than those observed for fine roots. The impact of nitrogen addition on HL, RB and RC significantly decreased with increasing experiment duration, amount of nitrogen addition and/or soil total phosphorus, conversely, its effect on HL and RL significantly intensified with soil total organic carbon. The effects of nitrogen and organic nutrient additions on RB exhibited significant positive and negative correlations, respectively, with their effects on HL in ectomycorrhizal hosts. No significant relationships were detected for arbuscular mycorrhizal hosts. Conclusion Our results demonstrate that plants preferentially employ root-dependent acquisition strategies under high inorganic nutrients but rely more on hypha-dependent strategies under abundant organic nutrients. This study underscores the necessity of integrating both roots and hyphae within current nutrient acquisition frameworks to evaluate the effects of global changes on carbon and nitrogen cycling. mycorrhizal nitrogen nutrient acquisition phosphorus soil organic matter Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 1 Introduction Plants obtain inorganic nutrients primarily through direct uptake by their fine roots, while simultaneous releasing root exudates, such as organic acids and enzymes, that stimulate microbial priming effects, thereby promoting the degradation of leaf litter and dead fine roots, the mineralization of soil organic matter, and the weathering of soil minerals to induce the release of associated nutrients such as nitrogen and phosphorus (Jones et al. 2004 ; Lambers et al. 2008 ; Vives-Peris et al. 2020 ). Moreover, by allocating carbon to symbiotic mycorrhizal fungi, plants enhance their access to both inorganic and organic forms of nitrogen and phosphorus, which these fungi mobilize with their hyphae from solid inorganic and organic materials or take up from soil solution (Wang et al. 2019 ; Jörgensen et al. 2024 ). For instance, the hyphal exudates of mycorrhizal fungi can hydrolyze organic nitrogen and phosphorus through their enzymatic activities, such as proteases and phosphatases, increasing nutrient transfer into the host plant roots (Finlay et al. 2020 ; Luthfiana et al. 2021 ). Consequently, plant nutrient acquisition strategies for exploiting soil resources can be categorized into two main pathways: fine-root proliferation, involving carbon investment into fine-root system expansion to directly absorb mineralized nutrients, and mycorrhizal symbiosis, characterized by carbon allocation to fungal hyphae that acquire nitrogen and phosphorus from inorganic and organic sources (Chari et al. 2024; Guo et al. 2025 ). Typically, plants employ a combination of these root and hyphal mechanisms to optimize nutrient uptake; however, the hyphae of certain fungal species have access to a broader reservoir of energy, nitrogen, and phosphorus compared to roots alone, e.g. via siderophores (Colpaert et al. 1992 ; Zou et al. 2019 ). The potential reduction in root nutrient-scavenging capacity resulting from root growth may theoretically be compensated by increased carbon investment in hyphal nutrient-acquisition strategies (Miller et al. 1995 ; Li et al. 2025 ). Therefore, considering the pivotal roles of both roots and hyphae in nutrient uptake, an integrated and concurrent examination of the plasticity of root and hyphal traits is essential to deepen our understanding of plant resource acquisition strategies under the influence of global environmental changes (Ma et al. 2021 ; Wang et al. 2021 ). Atmospheric nitrogen deposition, a critical component of global environmental change, exerts a substantial influence on the nutrient acquisition strategies of plants within terrestrial ecosystems (Wang et al. 2022 ; Jörgensen et al. 2024 ). Nevertheless, investigations into the effects of nitrogen availability on root and fungal hyphal growth have yielded heterogeneous outcomes, including increases, decreases and negligible alterations (Högberg et al. 2021 ; Wang et al. 2021 ; Jörgensen et al. 2024 ). These variable responses are attributed to the complex interactions between nitrogen addition and soil biotic and abiotic factors, such as soil pH, host carbon allocation and the duration of nitrogen addition (Wallander 1995 ; Lin et al. 2017 ; Wang et al. 2019 ; Ma et al. 2021 ; Marshall et al. 2023 ). Furthermore, distinct mycorrhizal symbioses, including arbuscular mycorrhizal and ectomycorrhizal associations, display divergent nutrient economy characteristics in plants inhabiting in terrestrial ecosystems (Chari et al. 2024; Chen et al. 2025 ). For example, plants associated with arbuscular mycorrhizae are generally more efficient in utilizing inorganic nutrients, whereas those forming ectomycorrhizal associations possess the capacity to access nitrogen and phosphorus from organic sources (Read 1991 ; Liu et al. 2018 ; Guo et al. 2025 ). Additionally, the inherent carbon utilization strategies of mycorrhizal fungi contribute to substantial variability in the responses of hyphae and roots to nitrogen deposition (Lin et al. 2017 ). Despite these insights, the precise direction and magnitude of fine-root and fungal hyphal responses to soil nitrogen availability remain inadequately understood. Elucidating these responses is essential for advancing our comprehension of plant nutrient acquisition strategies and for identifying the principal factors that regulate these dynamics (Wang et al. 2021 ). Prolonged nitrogen deposition has been shown to markedly decrease soil phosphorus concentrations by inhibiting soil acid phosphatase activity, thereby reducing the availability of soil phosphorus (Bowman et al. 2008 ; Zhang et al. 2024 ). Consequently, phosphorus increasingly becomes a limiting nutrient for the growth of mycorrhizal hyphae as nitrogen availability rises (Wang et al. 2019 ). The observed shift in the ectomycorrhizal fungal communities, characterized by a decline in hyphal abundance under elevated soil nitrogen conditions, likely diminishes the capacity for phosphorus acquisition. This phenomenon may represent a key mechanism underlying nitrogen deposition-induced phosphorus limitation in numerous forest ecosystems (Wang et al. 2022 ). Conversely, enhanced soil carbon availability — resulting from increased leaf litterfall and fine-root biomass due to alleviated phosphorus constraints — may partly explain the positive impact of phosphorus supplementation on mycorrhizal hyphae (Liu et al. 2012 ; Li et al. 2015 ). However, given the substantial variability in climate, soil characteristics, and phosphorus availability across tropical to boreal biomes, comprehensive global-scale investigations are necessary to elucidate the effects of soil phosphorus on root and fungal hyphal dynamics (Huang et al. 2016 ; Wang et al. 2019 ). Global warming deeply impacts soil organic carbon dynamics by affecting the decomposition and accumulation of charcoal deposited by wildfire (Sun et al. 2015 ; Bryanin and Makoto 2017 ). Moreover, the application of wood ash or of organic matter is a prevalent practice in forest management (Hagerberg and Wallander 2002 ; Hammer et al. 2011 ). The advantageous effects of soil organic matter are associated with enhancements in physical soil properties, including increased porosity, improved water retention, and synergistic microbial activity, as well as a reduction in mechanical resistance to root and hyphal penetration (Ryan et al. 1994 ; Joner and Jakobsen 1995 ). Ectomycorrhizal fungi, in particular, allocate substantial carbon to synthesize a variety of hydrolytic and oxidative enzymes that facilitate the degradation of carbon-containing compounds and the mobilization nutrients from soil organic matter (Courty et al. 2010 ; Chen et al. 2025 ). Consequently, it is hypothesized that mycorrhizal hyphae exhibit a more pronounced responses to the addition of organic matter compared to roots. However, the mechanisms governing the colonization of added wood ash or of added organic matter by mycorrhizal fungi remain poorly understood (Joner and Jakobsen 1995 ; Labidi et al. 2007 ; Majdi et al. 2008 ). Collectively, advancing our comprehension on the integrated responses of roots and hyphae to alterations in soil organic matter is essential for accurately predicting the effects of climate warming on plant nutrient acquisition strategies. Currently, the absence of concurrent investigations on fine roots and fungal hyphae limits our insight into how mycorrhizal plants adapt their nutrient foraging strategies in response to global climate change (Wang et al. 2021 ). To address these knowledge gaps, we conducted a comprehensive global meta-analysis that simultaneously integrated root and hyphal data derived from experiments involving nitrogen, phosphorus and organic matter additions. We hypothesized that: (1) roots would exhibit pronounced responses to inorganic nutrient additions, whereas hyphae would demonstrate stronger responses to organic nutrient additions; (2) the responses of roots and hyphae to nutrient additions would be modulated by abiotic and/or biotic factors, including fertilization regimes, soil properties and mycorrhizal types. More specifically, we anticipated that the effects of nutrient additions on roots and hyphae would diminish with increasing experiment duration and higher levels of soil nitrogen, phosphorus and organic carbon. 2 Materials and methods 2.1 Data collection We conducted a comprehensive literature search utilizing the Web of Science and China National Knowledge Infrastructure (CNKI) to identify studies that concurrently assessed the responses of roots and hyphae to nitrogen, phosphorus, nitrogen-phosphorus combination and organic nutrient additions. The keywords were “root response” or “mycelia response” or “plant and fungal response” or “root-fungal dynamics” or “mycorrhizal fungal growth” or “mycorrhizal fungal mycelium” or “fungal hyphae” or “extramatrical mycelia” or “nutrient addition” or “nitrogen supply” or “nitrogen enrichment” or “nitrogen input” or “nitrogen deposition” or “nitrogen fertilization” or “nitrogen amendment” or “phosphorus fertilization” or “phosphorus addition”, or “phosphorus supply” or “wood ash” or “organic addition” or “organic fertilization” or “organic amendments” or “organic nutrient patches” or “organic matter” or “compost addition”. In this study, we selected publications based on the following criteria: (1) experiments must have been conducted in terrestrial ecosystems (e.g. forests, meadows, shrubland and cropland; Table S1), including soil-based incubation studies conducted indoors; (2) experiments must encompass both control and nutrient addition treatments, as well as both root and hyphal observations under the same abiotic and biotic conditions; (3) the average values, standard deviation/standard errors and numbers of samples are available or can be calculated; (4) hyphae from ingrowth bags and soil cores are estimated by measuring the hyphal biomass using biomarkers (e.g. ergosterol and phospholipid fatty acid) and the hyphal length using non-biomarkers (e.g. gridline intersection and agar film). It is undeniable that hyphae in ingrowth bags and soil cores also can be hyphae from non-mycorrhizal fungi, such as saprotrophic fungi (Chen et al. 2018 ; Wang et al. 2021 ), however, as demonstrated by DNA analyses, the majority of hyphae is of mycorrhizal origin (Wallander et al. 2010). When several mycorrhizal fungal species or host plant species were reported in a study, we considered them as independent data points. If parameters were measured several times in a study, the last sampling data was used in this meta-analysis. After applying the selection criteria, a total of 738 observation pairs concerning root and hyphae were compiled, encompassing five variables: hyphal length (HL), hyphal biomass (HB), root length (RL), root biomass (RB) and root colonization (RC). In arbuscular mycorrhizal host plants, root colonization was quantified as the proportion of root length occupied by mycorrhizal fungi. Conversely, in ectomycorrhizal host plants, root colonization was determined by the percentage of root tips colonized by mycorrhizal fungi (Chen et al. 2016 ; Wang 2021 ). Totally, these observations examined the effects of nitrogen addition (301 observations), phosphorus addition (193 observations), nitrogen-phosphorus combination addition (83 observations) and organic nutrient addition (161 observations) on the five variables across 43 distinct experiments. In this study, we collected the mean, sample size and standard deviation of the treatment and control data from each study. We obtained the data directly from tables or extracted them from figures using GetData Graph Digitizer 2.24 ( http://getdata-graph-digitizer.com ). We recorded various site-specific characteristics, including geographic coordinates (longitude and latitude), climatic variables (mean annual temperature (MAT) and aridity index (AI)), soil properties (soil total nitrogen (TN), soil total phosphorus (TP), soil total organic carbon (TOC) and soil pH), fertilization regimes (experiment duration (DU) and amount of nutrient addition (AM)), measurement and sampling methods of root (RM) and hyphal traits (HM), and mycorrhizal types of hosts (MT). Furthermore, we have divided mycorrhizal associations into two categories: arbuscular mycorrhizal hosts, which include trees, grasses and shrubs, and ectomycorrhizal hosts, which include trees (Table S1). We acquired MAT from the WordClim global climate layers, with a spatial resolution of about 1 km ( https://www.worldclim.org ), with a spatial resolution of 5° longitude by 3.75° latitude (Dentener 2006 ). We extracted AI from the Global Aridity Index and the Potential Evapotranspiration Database - Version 3, with a spatial resolution of 30 arc-seconds (Zomer et al. 2022 ). For edaphic factors, we extracted TOC, TN, TP, pH at 0–30 cm soil depth from the global soil dataset for use in Earth system models, with a 1 km spatial resolution (Shangguan et al. 2014 ). 2.2 Statistical analysis In this study, we selected the natural log-transformed response ratio (RR) to calculate the “effect size” of different treatments on hyphal and root traits (Hedges et al. 1999 ). We defined RR as the ratio of the value of a parameter in the treatment group to that in the control group. We employed a random effects model to calculate RR and to generate confidence intervals (CI) with the restricted maximum likelihood method using the rma.mv function in the metafor package version 4.8-0 (Viechtbauer 2010 ). If the 95% CI did not overlap with zero, the effects of treatments on the response variables were considered statistically significant at P < 0.05. We explored the possibility of publication bias using Egger’s regression test ( Z - and P -values are given), trim and fill model and bootstrap approach (Egger et al. 1997 ; Veroniki et al. 2016 ; Rossetti et al. 2017 ) to examine the validity of the results of our meta-analysis. Moreover, RR and its CI were transformed back to the percentage change (% Change) for ease of interpretation. We employed linear regression and generalized additive models to evaluate the relationships between the effects of nitrogen/organic additions on roots and their effects on hyphae for each mycorrhizal type and for the aggregated dataset, respectively. Moreover, we used linear regression analysis to highlight the relationships of effect sizes of nitrogen/organic nutrient additions on roots and hyphae in relation to various soil and experimental variables. MetaForest, a random forest-based algorithm, takes into account different weighting between experiments, incorporates several different predictors and their interactions, and considers the non-linear relationship between moderators and the predicted variables (van Lissa 2020 ). In this study, we used MetaForest analysis to determine the relative importance of climate, soil properties, mycorrhizal types, root sampling methods, and fertilization regimes in modulating the responses of roots and hyphae to nutrient additions. The analysis was conducted using the MetaForest function in the MetaForest package version 0.1.4. Our analytical procedure initially involved assessing the convergence of the original model, followed by preselecting the moderators, tuning the model parameters, selecting the final model, and then verifying its convergence. We evaluated model performance using the retrodictive R 2 ( R 2 fit ) and the predictive R 2 metrics derived from cross-validation ( R 2 cv ) and out-of-bag testing ( R 2 oob ) (van Lissa 2020 ). 3 Results 3.1 Responses of hyphae and roots to nitrogen, phosphorus and organic nutrient additions Nitrogen addition resulted in a significant increase in hyphal length (26%), root biomass (34%) and root length (31%), while concurrently causing a significant reduction in root colonization (-20%). Phosphorus addition led to notable enhancements in root biomass (27%) and root length (36%), but did not significantly influence hyphal biomass, hyphal length or root colonization. Organic nutrient addition significantly increased hyphal biomass (63%) and length (268%), as well as root biomass (87%) and root length (24%), without affecting root colonization (Fig. 1 ). Moreover, the combined application of nitrogen and phosphorus addition did not produce significant changes in any hyphal or root traits (Fig. 1 ). Notably, roots and hyphae associated with ectomycorrhizal hosts showed more pronounced responses to nutrient additions compared to those of arbuscular mycorrhizal hosts (Figure S1). Comprehensive analyses employing Egger’s regression test, the trim and fill method and bootstrap approaches confirmed that the directionality and statistical significance of effect sizes related to nitrogen, phosphorus, combined nitrogen and phosphorus, and organic nutrient addition remained robust despite potential publication bias (Table S2). Linear regression analyses revealed a positive linear relationship between the effects of nitrogen addition on root biomass and ectomycorrhizal hyphal length, whereas a negative linear relationship was observed between the effects of organic nutrient addition on root biomass and ectomycorrhizal hyphal length (Fig. 2 ). In contrast, no significant relationships were detected for arbuscular mycorrhizal hosts (Fig. 2 ). When data were aggregated, the linear and nonlinear relationships emerged between the effects of nitrogen and organic nutrient additions on root biomass and their corresponding effects on hyphal length, respectively (Fig. 2 ). 3.2 Key factors influencing hyphae and roots response to nitrogen and organic nutrient additions MetaForest modeling accounted for 45%, 71%, 83% and 51% of the variance in the effects of nitrogen addition on hyphal length, root biomass, root length and root colonization, respectively, identifying fertilization regime, climatic variables, and soil properties as important moderators (Table 1 ; Fig. 3 ). Specifically, experiment duration and soil total phosphorus were the two most influential factors affecting the impact of nitrogen addition on hyphal length and root biomass (Fig. 3 a, b), whereas soil total organic carbon and mean annual temperature, as well as amount of nitrogen addition and experiment duration were critical determinants for root length and root colonization, respectively (Fig. 3 c, d). The effects of nitrogen addition on hyphal length, root biomass and root colonization exhibited significant declines with increasing experiment duration/amount of nitrogen addition (Fig. 4 a-c). Additionally, nitrogen-induced effects on root length and root colonization significantly decreased with mean annual temperature and latitude, respectively (Fig. 4 d, f), while effects on root length significantly increased with latitude (Fig. 4 e). Effects of nitrogen addition on hyphal length, root biomass and root colonization were negatively correlated with soil total phosphorus or soil pH (Fig. 5 a, b, d, h), whereas root length and hyphal length significantly increased with soil total phosphorus, soil total organic carbon and soil pH (Fig. 5 c, e-g). Regarding organic nutrient addition, MetaForest models explained 53% and 80% of the variation in their effects on organic nutrient addition on hyphal biomass and root biomass, respectively, with mycorrhizal type and soil properties (such as soil total nitrogen) serving as key moderators (Table 1 ; Figure S2a, b). Moreover, mycorrhizal type and soil total organic carbon emerged as the first influential factors governing the effects of phosphorus addition on root biomass and root length (Figure S2c, d). Table 1 Performance of MetaForest models in predicting effects of nitrogen (N), phosphorus (P), N-P combination (NP) and organic nutrient (O) additions on hyphal biomass (HB), hyphal length (HL), root biomass (RB), root length (RL) and root colonization (RC). Model performance is evaluated using the retrodictive R 2 ( R 2 fit ), and the predictive R 2 values obtained during cross validation ( R 2 cv ) and out-of-bag tests ( R 2 oob ). Non-negative R 2 oob values indicate no serious model overfitting. R 2 fit R 2 cv R 2 oob N P NP O N P NP O N P NP O HB - - - 0.53 - - - 0.29 - - - 0.03 HL 0.45 - - - 0.32 - - - 0.12 - - - RB 0.71 0.23 - 0.80 0.62 0.71 - 0.78 0.46 0.50 - 0.59 RL 0.83 0.06 - - 0.38 0.32 - - 0.10 0.10 - - RC 0.51 - - - 0.36 - - - 0.20 - - - Table S1 The reference list (the details on references are provided in the data sources section at the conclusion of the text) regarding the classifications of terrestrial ecosystems, the types of mycorrhizae and the categories of nutrient additions utilized in this study. References Inorganic nutrient addition Organic nutrient addition (O) Nitrogen (N) Phosphorus (P) and wood ash (P) N-P combination (NP) Grasses/Meadows, with AM Yang Y, Zhang J, He J et al (2023) NH 4 NO 3 Ca(H 2 PO 4 ) 2 NH 4 NO 3 , Ca(H 2 PO 4 ) 2 Antoninka A, Reich PB and Johnson NC (2011) NH 4 NO 3 Liu Y, Shi G, Mao L et al (2012) (NH 4 ) 2 HPO 4 Muneer MA, Wang P, Zaib-un-Nisa et al (2020) NH 4 Cl Muneer MA, Wang P, Zhang J et al (2020) NH 4 Cl Shrubs/Shrubland, with AM Rillig MC and Allen MF (1998) CaNO 3 , NH 4 NO 3 Pan S, Wang Y, Qiu Y et al (2020) NH 4 NO 3 Trees/Crops/Forest, with AM Yan G, Zhou M, Wang M et al (2019) NH 4 NO 3 Qu Z, Zhu L, Jiang Q et al (2024) NaH 2 PO 4 ·2H 2 O Chen W, Koide RT, Adams TS et al (2016) Leaf Camenzind T, Homeier J, Dietrich K et al (2016) CO(NH 2 ) 2 NaH 2 PO 4 ·2H 2 O CO(NH 2 ) 2 , NaH 2 PO 4 ·2H 2 O Liu B, Li L, Rengel Z et al (2019) CO(NH 2 ) 2 NaH 2 PO 4 Zheng C, Chai M, Jiang S et al (2015) KH 2 PO 4 Li L, McCormack ML, Chen F et al (2019) NH 4 NO 3 NaH 2 PO 4 NH 4 NO 3 , NaH 2 PO 4 Cheng L, Chen W, Adams TS et al (2016) NH 4 NO 3 , NaH 2 PO 4 Leaf Bethlenfalvay GJ (1983) Ca 10 (PO 4 ) 6 (OH) 2 Frey JE and Ellis JR (1997) KH 2 PO 4 , NH 4 NO 3 Staddon PL, Jakobsen I and Blum H (2004) NH 4 NO 3 Hodge A and Fitter AH (2010) NA Leaf Cao C (2021) NH 4 NO 3 Zhang F (2019) CO(NH 2 ) 2 Chen W, Koide RT and Eissenstat DM (2018) Leaf Trees/Forest, with ECM : Alberton O and Kuyper TW (2009) NA Yan G, Zhou M, Wang M et al (2019) NH 4 NO 3 Zhang J, Lin G and Zeng D-H (2024) CO(NH 2 ) 2 Chen W, Koide RT, Adams TS et al (2016) Leaf Ekblad A, Wallander H, Carlsson R et al (1995) NH 4 NO 3 KH 2 PO 4 , K 2 HPO 4 , NaH 2 PO 4 , Na 2 HPO 4 NaH 2 PO 4 , KH 2 PO 4 , K 2 HPO 4 , NaH 2 PO 4 , Na 2 HPO 4 Kårén O and Nylund JE (1996) (NH 4 ) 2 SO 3 Leppalammi-Kujansuu J, Ostonen I, Stromgren M et al (2013) NH 4 NO 3 Weigt RB, Raidl S, Verma R et al (2011) NH 4 NO 3 Wiemken V, Ineichen K and Boller T (2001) NH 4 NO 3 Cheng L, Chen W, Adams TS et al (2016) NH 4 NO 3 , KH 2 PO 4 Leaf Majdi H, Truus L, Johansson U et al ( 2008 ) Wood ash granules Jones MD, Durall DM and Tinker PB (1990) KH 2 PO 4 Pampolina NM, Dell B and Malajczuk N (2002) Ca(H₂PO₄) 2 ·CaHPO₄ Treseder KK, Turner KM and Mack MC (2007) NH 4 NO 3 Bakker MR, Jolicoeur E, Trichet P et al (2009) NA Naples BK and Fisk MC (2010) NH 4 NO 3 NaH 2 PO 4 Rasheed MU, Kasurinen A, Kivimäenpää M et al (2017) Yara Peatcare TM Zhu X, Zhang Z, Wang Q et al (2022) NH 4 NO 3 Wang C, Brunner I, Guo W et al ( 2021 ) NH 4 NO 3 He Y (2021) Ca(H 2 PO 4 ) 2 ·H 2 O Wang G ( 2021 ) Osmocote Chicken manure Forsmark B, Nordin A, Rosenstock NP et al (2021) NH 4 NO 3 Du Z, Wang W, Zeng W et al (2014) CO(NH 2 ) 2 Chen W, Koide RT and Eissenstat DM (2018) Leaf Table S2 Results of testing publication bias for statistically significant response variables for nitrogen (N), phosphorus (P), N-P combination (NP) and organic nutrient (O) additions on hyphal biomass (HB), hyphal length (HL), root biomass (RB), root length (RL) and root colonization (RC) using Egger’s regression test ( Z - and P -values are given), trim and fill model and bootstrap approach (5% and 95% confidence intervals [CI] of effect sizes are given). Egger’s regression Trim and fill model Bootstrap approach Original model Treatments Variables Z P 5%CI 95%CI 5%CI 95%CI 5%CI 95%CI N HB -1.213 0.225 -0.063 0.252 -0.118 0.112 -0.149 0.148 HL 1.486 0.137 0.152 0.403 0.181 0.376 0.152 0.403 RB -1.874 0.060 0.443 0.770 0.174 0.417 0.130 0.457 RL 2.537 0.011 -0.010 0.293 0.150 0.391 0.122 0.413 RC -3.261 0.001 -0.264 -0.025 -0.302 -0.137 -0.329 -0.120 P HB 3.019 0.003 -0.221 0.152 -0.166 0.111 -0.221 0.152 HL 1.180 0.238 -0.034 0.307 -0.011 0.271 -0.034 0.307 RB -0.175 0.861 0.161 0.419 0.136 0.357 0.113 0.371 RL -0.440 0.660 0.317 0.724 0.175 0.461 0.123 0.489 RC -2.623 0.009 -0.358 0.014 -0.324 -0.020 -0.358 0.014 NP HB -0.937 0.349 -0.287 0.159 -0.226 0.106 -0.287 0.159 HL -1.725 0.085 -0.529 0.194 -0.491 0.131 -0.529 0.194 RB -1.117 0.264 -0.315 0.286 -0.267 0.278 -0.315 0.286 RL -1.651 0.099 -0.148 0.273 -0.239 0.070 -0.299 0.084 RC 0.974 0.330 -0.283 0.054 -0.117 0.114 -0.158 0.154 O HB 0.889 0.374 0.196 0.684 0.214 0.689 0.249 0.723 HL 0.176 0.860 1.244 1.361 1.267 1.370 1.244 1.361 RB -1.713 0.087 0.387 1.193 0.320 0.952 0.220 1.034 RL 0.865 0.387 -0.065 0.267 0.102 0.333 0.068 0.367 RC -0.342 0.732 -0.051 0.142 -0.050 0.137 -0.056 0.138 4 Discussion 4.1 Variations in nutrient acquisition of roots and hyphae in response to inorganic and organic nutrient additions This study revealed that roots exhibited more pronounced positive responses to the addition of inorganic nutrients, specifically nitrogen and phosphorus compared to hyphae (Fig. 1 ), supporting our first hypothesis. Typically, plants tend to prioritize root proliferation under conditions of abundant inorganic nutrient availability, thereby employing an inorganic nutrient acquisition strategy (Mohan et al. 2014 ; Chari et al. 2024; Wang et al. 2025 ). Although hyphal length significantly increased following nitrogen addition, a concurrent decrease in root colonization was observed, suggesting that plants may reduce their reliance on mycorrhizal associations while enhancing direct nitrogen uptake through roots under elevated nitrogen inputs (Fig. 6 ). However, in comparison to roots, mycorrhizal fungi require a greater carbon investment due to their rapid hyphal turnover rates (Jakobsen and Rosendahl 1990 ; Ma et al. 2021 ). Consequently, plants reduce the carbon maintenance costs associated with nitrogen acquisition by allocating more carbon to fine roots and less to mycorrhizal hyphae as soil nitrogen availability increases, particularly in regions characterized by lower temperatures and higher latitudes (Fig. 4 d-f). Furthermore, phosphorus addition significantly increased root length and root biomass but did not affect hyphal length, hyphal biomass or root colonization (Fig. 1 ). Previous studies have demonstrated that plants can alter their root morphology to optimize inorganic phosphorus acquisition by producing more finely branched roots and elongating root hairs (Lambers et al. 2008 ; White et al. 2013 ; Péret et al. 2014 ). These results suggest that plants with an adequate supply of inorganic phosphorus prefer to use their roots for phosphorus uptake, as they exhibit higher uptake efficiency and physiological performance compared to phosphorus-deficient conditions, where mycorrhizal symbiosis plays a more important role (Brown et al. 2013 ; Wang et al. 2025 ). Consequently, plants employ root-dependent nutrient strategies in response to elevated inorganic nitrogen and phosphorus availability (Fig. 6 ). In this study, we observed an increase in root biomass and root length after the addition of organic nutrients, suggesting that plants may release root exudates that stimulate microbial priming effects, thereby promoting the weathering of soil minerals and the mineralization of soil organic matter, and thus also the release of the associated nutrients (Li et al. 2025 ). Nonetheless, the overall root responses to organic addition were relatively modest compared to the pronounced positive effects observed on fungal hyphae (Fig. 1 ), again confirming our first hypothesis. The enhancement of fungal hyphae due to organic nutrient addition is attributed to the beneficial impacts of increased organic matter on soil water status, soil structure and synergistic microbial activity, and the reduction of mechanical resistance to hyphal penetration within the soil (Ryan et al. 1994 ; Joner and Jakobsen 1995 ; Gryndler et al. 1998 ; Hammer et al. 2011 ). As expected in our second hypothesis, our findings indicate that soil total organic carbon is a critical factor modulating the effects of nitrogen addition on both hyphal length and root length, with positive correlations observed between these variables and soil total organic carbon (Figs. 3 a, c and 5 e, f). Soil organic carbon contributes to the stabilization of soil structure and enhances soil cation exchange capacity, both of which are essential for soil nutrient retention within the soil (Chen et al. 2023 ). Consequently, soils with a higher soil organic carbon content are able to retain larger amounts of plant-available nitrogen under the same nitrogen additions, thereby eliciting a more pronounced root response to the nitrogen supplementation. Moreover, the accelerated growth observed in soils enriched with the higher organic matter may confer a selective advantage to fungal hyphae in forest ecosystems frequently subjected to disturbance such as wildfires, facilitating enhanced nutrient acquisition from post-fire ash deposits (Majdi et al. 2008 ). The decomposition of organic matter in close proximity to the mycorrhizal hyphae provides a rich source of accessible carbon and energy (Gryndler et al. 1998 ). Accordingly, plants may preferentially adopt an organic nutrient acquisition strategy, wherein symbiotic associations promote microbial mineralization of nitrogen and phosphorus under conditions characterized by abundant organic nutrient availability (Fig. 6 ). 4.2 Factors influencing root and hyphal responses to inorganic and organic nutrient additions In this meta-analysis, a positive correlation was identified between the effects of nitrogen addition on root biomass and those on hyphal length in ectomycorrhizal hosts (Fig. 2 a). This associations is consistent with the premise that root and hyphal development are interconnected, as hyphal growth is driven by carbon allocation from fine roots, thereby fostering a positive relationship between root formation and fungal hyphae proliferation (Korkama et al. 2007 ). Elevated nitrogen deposition mitigates nitrogen limitation within ecosystems, thereby enhancing plant productivity, which is indicative of a greater allocation of carbon belowground, specifically to roots and their symbiotic mycorrhizae (Schulte-Uebbing and De Vries 2017 ; Wang et al. 2024 ). Nonetheless, plants must strategically allocate carbon resources between root-mediated nutrient uptake and hyphal nutrient acquisition mechanisms under nutrient-limited conditions (Wang et al. 2025 ). A prominent illustration of this resource allocation trade-off is the antagonistic interaction observed between mycorrhizal symbioses and fine-root growth, attributable to their morphological similarities and potential functional redundancy (Brown et al. 2013 ; Bergmann et al. 2020 ; Ma et al. 2021 ). This antagonism is corroborated by the significantly negative correlation found between the effects of organic nutrient amendments on root biomass and hyphal length in ectomycorrhizal hosts (Fig. 2 b). Compared to roots, ectomycorrhizal fungi exhibit greater efficiency in extracting nutrients from soil organic matter (Brown et al. 2013 ; Li et al. 2025 ; Wang et al. 2025 ). Collectively, these findings suggest that ectomycorrhizal plants may intensify their symbiotic interactions with mycorrhizal fungi, relying less on the proliferation of fine roots to optimize the acquisition of organic nutrients following organic nutrient amendments. In the present study, no significant correlation was observed between the effects of organic nutrient amendments on root biomass and their effects on hyphal length in arbuscular mycorrhizal hosts (Fig. 2 b). These results suggest that the influence of organic nutrient additions on root and hyphae development depends on the type of mycorrhiza (Figures S1, S2). The divergent nutrient acquisition strategies exhibited by arbuscular mycorrhizal and ectomycorrhizal hosts in response to organic nutrient amendments may be attributed to variations in environmental conditions and/or differential capacities to access nutrients from soil organic matter (Chari et al. 2024; Guo et al. 2025 ). Unlike arbuscular mycorrhizal hosts, which are predominantly distributed across tropical and subtropical regions, ectomycorrhizal hosts primarily inhabit temperate and boreal zones characterized by soils rich in organic matter but typically deficient in inorganic nitrogen (Reich and Oleksyn 2004 ; Ma et al. 2021 ). Consequently, ectomycorrhizal fungi generally exhibit superior capabilities in mineralizing soil organic matter compared to arbuscular mycorrhizal fungi (Rosling et al. 2016 ; Chen et al. 2018 ). The findings of this study underscore that the distinct responses of roots and hyphae to organic nutrient amendments, differentiated by mycorrhizal types, can substantially affect resource acquisition. Nonetheless, further research is required to elucidate the underlying mechanisms driving these observed patterns on arbuscular mycorrhizal versus ectomycorrhizal host plants (Fig. 6 ). The MetaForest analysis identified fertilization regime as the primary determinant influencing the impact of nitrogen addition on root and hyphal growth (Fig. 3 ), being supported by our second hypothesis. Specifically, short-term nitrogen supplementation appears to promote the development of roots and hyphae; however, this beneficial effect diminishes with extended nitrogen application, particularly beyond a decade. This pattern can be elucidated by several mechanisms. Initially, nitrogen addition enhances plant growth and increases carbon allocation to belowground structures, but these effects decline as the duration of nitrogen exposure lengthens (Li et al. 2016 ; Song et al. 2019 ). This observation aligns with findings from a recent meta-analysis demonstrating a temporal decrease in nitrogen-induced fungal biomass enhancement (Ma et al. 2021 ). Furthermore, prolonged nitrogen addition may lead to soil acidification, which adversely affects root and microbial proliferation, thereby counteracting the initial positive effects of nitrogen enrichment (Sterkenburg et al. 2015 ). This is corroborated by the significant positive correlation observed between nitrogen addition effects on root length and soil pH (Fig. 5 g). Moreover, our observations indicate that root colonization markedly declines as the amount of nitrogen supplementation increases (Fig. 4 c), implying that plants predominantly acquire nutrients through fine roots rather than mycorrhizal fungi when nitrogen availability in the soil is elevated (Ma et al. 2021 ). Taken together, these convergent lines of evidence underscore fertilization regime, such as experiment duration and amount of nutrient addition, as a critical factor modulating both the direction and magnitude of nitrogen addition effects on roots and hyphae. This study identified soil total phosphorus as a key factor influencing the variability in hyphal and root responses to nitrogen amendment (Fig. 5 a-d). Nitrogen addition promoted hyphal proliferation, as evidenced by increased hyphal length and root colonization under conditions of low phosphorus availability (Fig. 5 a, d). Generally, the abundance of root-external hyphae is positively associated with phosphorus acquisition efficiency in mycorrhizal plants (de Miranda and Harris 1994 ; Sbrana et al. 2022 ). Consequently, the initial enhancement of hyphal length and root colonization partially mitigates phosphorus limitation for plant growth as soil nitrogen availability rises. Negative growth responses in mycorrhizal hyphae may occur when carbon-limited fungi must allocate a greater proportion of host-derived carbon to excessive phosphorus assimilation rather than to growth. Conversely, nitrogen addition exerts more pronounced positive effects on root growth, as indicated by increased root length in areas with higher phosphorus concentrations (Fig. 5 c). Collectively, these results suggest a shift in plant nutrient acquisition strategies from reliance on mycorrhizal associations toward greater dependence on fine roots (e.g., increased root length observed in this study) with increasing soil phosphorus levels. Thus, plants appear to strategically allocate limited carbon resources between root and mycorrhizal traits, balancing these two principal pathways for phosphorus uptake (Wang et al. 2025 ). However, this proposed mechanism could not be confirmed through combined nitrogen and phosphorus additions due to insufficient paired data in the present study. Therefore, future research should prioritize investigating plant mycorrhizal symbioses and fine-root traits in response to simultaneous nitrogen and phosphorus supplementation. 5. Conclusions Our meta-analysis elucidated the complex responses of roots and fungal hyphae to nitrogen, phosphorus, and organic nutrient additions, indicating shifts in plant nutrient acquisition strategies in relation to the availability of inorganic and/or organic soil nutrients. The results demonstrated that roots possess advantages over hyphae in terms of root length and biomass, thereby exhibiting greater adaptability to inorganic nutrient (nitrogen and phosphorus) supplementation. Conversely, fungal hyphae, particularly those associated with ectomycorrhizal hosts, showed enhanced growth and distinct advantages relative to roots following organic nutrient additions. Additionally, a significant positive correlation was observed between the effect sizes on root biomass and ectomycorrhizal hyphal length under nitrogen addition, whereas a negative correlation was found under organic addition. Moreover, our findings suggest that fertilizer regimes (e.g. experiment duration and amount of nutrient addition), climatic factors (e.g. mean annual temperature), soil characteristics (e.g. pH, phosphorus, and organic carbon concentrations), and mycorrhizal types collectively influence the responses of roots and hyphae to nitrogen and organic amendments. These divergent responses underscore the necessity of integrating both root and hyphal dynamics within nutrient acquisition frameworks to accurately evaluate and predict the consequences of global environmental changes on carbon, nitrogen, and phosphorus cycling. Declarations Declaration of interests The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgments This work was financially supported by the Natural Science Foundation of China (grant number 42171051). 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06:38:40","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7788604/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7788604/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11104-026-08389-8","type":"published","date":"2026-02-20T15:59:19+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":94146306,"identity":"3882b9a5-f96f-4168-8f96-7b82e22d97f4","added_by":"auto","created_at":"2025-10-22 21:36:52","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1343245,"visible":true,"origin":"","legend":"","description":"","filename":"FiguresandtablesS2025106.docx","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/21160273aac1b74ed8844823.docx"},{"id":94146301,"identity":"ab34117b-fff5-4687-8f47-b5a73e5cf46a","added_by":"auto","created_at":"2025-10-22 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21:36:52","extension":"xml","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":194734,"visible":true,"origin":"","legend":"","description":"","filename":"PLSOD25038150structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/d0546b70cfcb99df7ce8acf8.xml"},{"id":94146473,"identity":"712d8e0b-5004-425a-bf5e-95113718d55a","added_by":"auto","created_at":"2025-10-22 21:44:52","extension":"html","order_by":24,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":202012,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/95d09a1f99b577b801b9d8a4.html"},{"id":94146300,"identity":"d68574f0-6466-4f41-9861-933ff7998c0d","added_by":"auto","created_at":"2025-10-22 21:36:52","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":489974,"visible":true,"origin":"","legend":"\u003cp\u003eResponses of hyphae and roots to nitrogen (N), phosphorus (P), N-P combination (NP) and organic nutrient (O) additions. HB, hyphal biomass; HL, hyphal length; RB, root biomass; RL, root length; RC, root colonization. The numbers on the top indicate the numbers of cases for hyphal and root traits. For ease of interpretation, the effect sizes (the natural log-transformed response ratio) and their corresponding confidence intervals were transformed back to the percentage change (% change). Error bars represent 95% confidence intervals. The effect of treatments is considered significant if the 95% confidence intervals of the effect size do not cover zero.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/fee0561c0ed8fad8d18b8011.png"},{"id":94146622,"identity":"308ed45f-040d-4410-abe7-9d9928d426e8","added_by":"auto","created_at":"2025-10-22 21:52:52","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":480374,"visible":true,"origin":"","legend":"\u003cp\u003eRelationships between the effect sizes of nitrogen (N; a) and organic (O; b) additions on root biomass (RB) and the effect sizes of N and O additions on hyphal length (HL) for arbuscular mycorrhiza (AM, indicated by the blue regression lines) and ectomycorrhiza (ECM, indicated by the red regression lines) hosts, as well as for the combined dataset (represented by the black regression lines).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/24925c349dea9ddf71a8af0b.png"},{"id":94146466,"identity":"4e3bea64-b606-4492-979b-871519ce9564","added_by":"auto","created_at":"2025-10-22 21:44:52","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1294823,"visible":true,"origin":"","legend":"\u003cp\u003eRelative importance of variables in predicting variation in the observed effects of nitrogen addition on hyphal length (HL, a), root biomass (RB, b) root length (RL, c) and root colonization (RC, d). Relative importance is quantified using MetaForest analyses. DU, experiment duration; TP, soil total phosphorus; AI, aridity index; pH, soil pH; TOC, soil total organic carbon; MAT, mean annual temperature; LA, latitude; TN, soil total nitrogen; AM, amount of nutrient addition; MT, mycorrhizal types; DE, sampling depth; RM, root estimation methods; HM, hyphae estimation methods; Fungi, fungi used at the community or individual level.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/6fb89131948f731b11f9d97f.png"},{"id":94146309,"identity":"54872515-7925-4c89-ac7c-529a473c0a54","added_by":"auto","created_at":"2025-10-22 21:36:52","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1001928,"visible":true,"origin":"","legend":"\u003cp\u003eVariations in the effects of nitrogen (N) addition on hyphal length (HL, a), root biomass (RB, b), root colonization (RC, c and f) and root length (RL, d and e) with experiment duration (a and b), amount of nitrogen addition (c), mean annual temperature (MAT, d) and/or latitude (LA, e and f). Dot sizes indicate the weights of the observations. Black lines represent the fitted linear regression lines and grey bands represent 95% confidence intervals.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/a0ad484daa8de6755588f980.png"},{"id":94146623,"identity":"b4209377-27f3-4a5c-bf99-e9bfed40faea","added_by":"auto","created_at":"2025-10-22 21:52:52","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":975843,"visible":true,"origin":"","legend":"\u003cp\u003eVariations in the effects of nitrogen (N) addition on hyphal length (HL, a and e), root biomass (RB, b), root length (RL, c, f and g) and root colonization (RC, d and h) with soil total phosphorus (TP, a-d), soil total organic carbon (TOC, e and f) and/or soil pH (pH, g and h). Dot sizes indicate the weights of the observations. Black lines represent the fitted linear regression lines and grey bands represent 95% confidence intervals.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/4430d2a2b78ab75dffbfa4fd.png"},{"id":94146470,"identity":"cdf609b3-4ffc-43aa-8671-8aaad6565fdb","added_by":"auto","created_at":"2025-10-22 21:44:52","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":694785,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of inorganic and organic amendments on nutrient acquisition strategies was examined by fine-root traits (root length, root biomass and root colonization) and mycorrhizal traits (hyphal length and root biomass). Blue and red arrows represent increases and decreases in the respective variables in response to inorganic (nitrogen and phosphorus; blue) and organic nutrient supplements (red), respectively. The magnitude of the arrows corresponds proportionally to the effect size of the nutrient treatments. The dashed grey arrows indicate that arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) host plants are hypothesized to utilize inorganic and organic nutrient acquisition strategies, respectively. Nevertheless, the precise effects of nutrient amendments on AM and ECM host plants remain unclear, owing to insufficient data within the scope of this meta-analysis.\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/4233262afefffe0ceb2bb8ae.png"},{"id":103252154,"identity":"60f6249c-ea1d-47bd-85da-a2e4dd141cc9","added_by":"auto","created_at":"2026-02-23 16:13:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":6249086,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/f18a0615-4888-4429-9adc-904c5aa852e7.pdf"},{"id":94146316,"identity":"3cf3a193-ca5a-4c23-86e3-515d8aab274c","added_by":"auto","created_at":"2025-10-22 21:36:52","extension":"png","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":631124,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/49e34f812a80fa1145bb930a.png"},{"id":94146319,"identity":"07d635bd-68c6-41b8-8d9f-4c97677d9cf9","added_by":"auto","created_at":"2025-10-22 21:36:52","extension":"png","order_by":14,"title":"","display":"","copyAsset":false,"role":"supplement","size":647321,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage8.png","url":"https://assets-eu.researchsquare.com/files/rs-7788604/v1/b4c3b6b829a36dd707cafb01.png"}],"financialInterests":"","formattedTitle":"Asymmetric responses of roots and hyphae to inorganic and organic nutrient additions: a global meta-analysis","fulltext":[{"header":"1 Introduction","content":"\u003cp\u003ePlants obtain inorganic nutrients primarily through direct uptake by their fine roots, while simultaneous releasing root exudates, such as organic acids and enzymes, that stimulate microbial priming effects, thereby promoting the degradation of leaf litter and dead fine roots, the mineralization of soil organic matter, and the weathering of soil minerals to induce the release of associated nutrients such as nitrogen and phosphorus (Jones et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Lambers et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Vives-Peris et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, by allocating carbon to symbiotic mycorrhizal fungi, plants enhance their access to both inorganic and organic forms of nitrogen and phosphorus, which these fungi mobilize with their hyphae from solid inorganic and organic materials or take up from soil solution (Wang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; J\u0026ouml;rgensen et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). For instance, the hyphal exudates of mycorrhizal fungi can hydrolyze organic nitrogen and phosphorus through their enzymatic activities, such as proteases and phosphatases, increasing nutrient transfer into the host plant roots (Finlay et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Luthfiana et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Consequently, plant nutrient acquisition strategies for exploiting soil resources can be categorized into two main pathways: fine-root proliferation, involving carbon investment into fine-root system expansion to directly absorb mineralized nutrients, and mycorrhizal symbiosis, characterized by carbon allocation to fungal hyphae that acquire nitrogen and phosphorus from inorganic and organic sources (Chari et al. 2024; Guo et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Typically, plants employ a combination of these root and hyphal mechanisms to optimize nutrient uptake; however, the hyphae of certain fungal species have access to a broader reservoir of energy, nitrogen, and phosphorus compared to roots alone, e.g. via siderophores (Colpaert et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Zou et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The potential reduction in root nutrient-scavenging capacity resulting from root growth may theoretically be compensated by increased carbon investment in hyphal nutrient-acquisition strategies (Miller et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Therefore, considering the pivotal roles of both roots and hyphae in nutrient uptake, an integrated and concurrent examination of the plasticity of root and hyphal traits is essential to deepen our understanding of plant resource acquisition strategies under the influence of global environmental changes (Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eAtmospheric nitrogen deposition, a critical component of global environmental change, exerts a substantial influence on the nutrient acquisition strategies of plants within terrestrial ecosystems (Wang et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; J\u0026ouml;rgensen et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Nevertheless, investigations into the effects of nitrogen availability on root and fungal hyphal growth have yielded heterogeneous outcomes, including increases, decreases and negligible alterations (H\u0026ouml;gberg et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; J\u0026ouml;rgensen et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These variable responses are attributed to the complex interactions between nitrogen addition and soil biotic and abiotic factors, such as soil pH, host carbon allocation and the duration of nitrogen addition (Wallander \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Lin et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Marshall et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Furthermore, distinct mycorrhizal symbioses, including arbuscular mycorrhizal and ectomycorrhizal associations, display divergent nutrient economy characteristics in plants inhabiting in terrestrial ecosystems (Chari et al. 2024; Chen et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For example, plants associated with arbuscular mycorrhizae are generally more efficient in utilizing inorganic nutrients, whereas those forming ectomycorrhizal associations possess the capacity to access nitrogen and phosphorus from organic sources (Read \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Additionally, the inherent carbon utilization strategies of mycorrhizal fungi contribute to substantial variability in the responses of hyphae and roots to nitrogen deposition (Lin et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Despite these insights, the precise direction and magnitude of fine-root and fungal hyphal responses to soil nitrogen availability remain inadequately understood. Elucidating these responses is essential for advancing our comprehension of plant nutrient acquisition strategies and for identifying the principal factors that regulate these dynamics (Wang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eProlonged nitrogen deposition has been shown to markedly decrease soil phosphorus concentrations by inhibiting soil acid phosphatase activity, thereby reducing the availability of soil phosphorus (Bowman et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Consequently, phosphorus increasingly becomes a limiting nutrient for the growth of mycorrhizal hyphae as nitrogen availability rises (Wang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The observed shift in the ectomycorrhizal fungal communities, characterized by a decline in hyphal abundance under elevated soil nitrogen conditions, likely diminishes the capacity for phosphorus acquisition. This phenomenon may represent a key mechanism underlying nitrogen deposition-induced phosphorus limitation in numerous forest ecosystems (Wang et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Conversely, enhanced soil carbon availability \u0026mdash; resulting from increased leaf litterfall and fine-root biomass due to alleviated phosphorus constraints \u0026mdash; may partly explain the positive impact of phosphorus supplementation on mycorrhizal hyphae (Liu et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). However, given the substantial variability in climate, soil characteristics, and phosphorus availability across tropical to boreal biomes, comprehensive global-scale investigations are necessary to elucidate the effects of soil phosphorus on root and fungal hyphal dynamics (Huang et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eGlobal warming deeply impacts soil organic carbon dynamics by affecting the decomposition and accumulation of charcoal deposited by wildfire (Sun et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Bryanin and Makoto \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Moreover, the application of wood ash or of organic matter is a prevalent practice in forest management (Hagerberg and Wallander \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Hammer et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). The advantageous effects of soil organic matter are associated with enhancements in physical soil properties, including increased porosity, improved water retention, and synergistic microbial activity, as well as a reduction in mechanical resistance to root and hyphal penetration (Ryan et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Joner and Jakobsen \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Ectomycorrhizal fungi, in particular, allocate substantial carbon to synthesize a variety of hydrolytic and oxidative enzymes that facilitate the degradation of carbon-containing compounds and the mobilization nutrients from soil organic matter (Courty et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Consequently, it is hypothesized that mycorrhizal hyphae exhibit a more pronounced responses to the addition of organic matter compared to roots. However, the mechanisms governing the colonization of added wood ash or of added organic matter by mycorrhizal fungi remain poorly understood (Joner and Jakobsen \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Labidi et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Majdi et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Collectively, advancing our comprehension on the integrated responses of roots and hyphae to alterations in soil organic matter is essential for accurately predicting the effects of climate warming on plant nutrient acquisition strategies.\u003c/p\u003e\u003cp\u003eCurrently, the absence of concurrent investigations on fine roots and fungal hyphae limits our insight into how mycorrhizal plants adapt their nutrient foraging strategies in response to global climate change (Wang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). To address these knowledge gaps, we conducted a comprehensive global meta-analysis that simultaneously integrated root and hyphal data derived from experiments involving nitrogen, phosphorus and organic matter additions. We hypothesized that: (1) roots would exhibit pronounced responses to inorganic nutrient additions, whereas hyphae would demonstrate stronger responses to organic nutrient additions; (2) the responses of roots and hyphae to nutrient additions would be modulated by abiotic and/or biotic factors, including fertilization regimes, soil properties and mycorrhizal types. More specifically, we anticipated that the effects of nutrient additions on roots and hyphae would diminish with increasing experiment duration and higher levels of soil nitrogen, phosphorus and organic carbon.\u003c/p\u003e"},{"header":"2 Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1 Data collection\u003c/h2\u003e\u003cp\u003eWe conducted a comprehensive literature search utilizing the Web of Science and China National Knowledge Infrastructure (CNKI) to identify studies that concurrently assessed the responses of roots and hyphae to nitrogen, phosphorus, nitrogen-phosphorus combination and organic nutrient additions. The keywords were \u0026ldquo;root response\u0026rdquo; or \u0026ldquo;mycelia response\u0026rdquo; or \u0026ldquo;plant and fungal response\u0026rdquo; or \u0026ldquo;root-fungal dynamics\u0026rdquo; or \u0026ldquo;mycorrhizal fungal growth\u0026rdquo; or \u0026ldquo;mycorrhizal fungal mycelium\u0026rdquo; or \u0026ldquo;fungal hyphae\u0026rdquo; or \u0026ldquo;extramatrical mycelia\u0026rdquo; or \u0026ldquo;nutrient addition\u0026rdquo; or \u0026ldquo;nitrogen supply\u0026rdquo; or \u0026ldquo;nitrogen enrichment\u0026rdquo; or \u0026ldquo;nitrogen input\u0026rdquo; or \u0026ldquo;nitrogen deposition\u0026rdquo; or \u0026ldquo;nitrogen fertilization\u0026rdquo; or \u0026ldquo;nitrogen amendment\u0026rdquo; or \u0026ldquo;phosphorus fertilization\u0026rdquo; or \u0026ldquo;phosphorus addition\u0026rdquo;, or \u0026ldquo;phosphorus supply\u0026rdquo; or \u0026ldquo;wood ash\u0026rdquo; or \u0026ldquo;organic addition\u0026rdquo; or \u0026ldquo;organic fertilization\u0026rdquo; or \u0026ldquo;organic amendments\u0026rdquo; or \u0026ldquo;organic nutrient patches\u0026rdquo; or \u0026ldquo;organic matter\u0026rdquo; or \u0026ldquo;compost addition\u0026rdquo;.\u003c/p\u003e\u003cp\u003eIn this study, we selected publications based on the following criteria: (1) experiments must have been conducted in terrestrial ecosystems (e.g. forests, meadows, shrubland and cropland; Table S1), including soil-based incubation studies conducted indoors; (2) experiments must encompass both control and nutrient addition treatments, as well as both root and hyphal observations under the same abiotic and biotic conditions; (3) the average values, standard deviation/standard errors and numbers of samples are available or can be calculated; (4) hyphae from ingrowth bags and soil cores are estimated by measuring the hyphal biomass using biomarkers (e.g. ergosterol and phospholipid fatty acid) and the hyphal length using non-biomarkers (e.g. gridline intersection and agar film). It is undeniable that hyphae in ingrowth bags and soil cores also can be hyphae from non-mycorrhizal fungi, such as saprotrophic fungi (Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), however, as demonstrated by DNA analyses, the majority of hyphae is of mycorrhizal origin (Wallander et al. 2010). When several mycorrhizal fungal species or host plant species were reported in a study, we considered them as independent data points. If parameters were measured several times in a study, the last sampling data was used in this meta-analysis. After applying the selection criteria, a total of 738 observation pairs concerning root and hyphae were compiled, encompassing five variables: hyphal length (HL), hyphal biomass (HB), root length (RL), root biomass (RB) and root colonization (RC). In arbuscular mycorrhizal host plants, root colonization was quantified as the proportion of root length occupied by mycorrhizal fungi. Conversely, in ectomycorrhizal host plants, root colonization was determined by the percentage of root tips colonized by mycorrhizal fungi (Chen et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Wang \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Totally, these observations examined the effects of nitrogen addition (301 observations), phosphorus addition (193 observations), nitrogen-phosphorus combination addition (83 observations) and organic nutrient addition (161 observations) on the five variables across 43 distinct experiments.\u003c/p\u003e\u003cp\u003eIn this study, we collected the mean, sample size and standard deviation of the treatment and control data from each study. We obtained the data directly from tables or extracted them from figures using GetData Graph Digitizer 2.24 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://getdata-graph-digitizer.com\u003c/span\u003e\u003cspan address=\"http://getdata-graph-digitizer.com\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). We recorded various site-specific characteristics, including geographic coordinates (longitude and latitude), climatic variables (mean annual temperature (MAT) and aridity index (AI)), soil properties (soil total nitrogen (TN), soil total phosphorus (TP), soil total organic carbon (TOC) and soil pH), fertilization regimes (experiment duration (DU) and amount of nutrient addition (AM)), measurement and sampling methods of root (RM) and hyphal traits (HM), and mycorrhizal types of hosts (MT). Furthermore, we have divided mycorrhizal associations into two categories: arbuscular mycorrhizal hosts, which include trees, grasses and shrubs, and ectomycorrhizal hosts, which include trees (Table S1). We acquired MAT from the WordClim global climate layers, with a spatial resolution of about 1 km (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.worldclim.org\u003c/span\u003e\u003cspan address=\"https://www.worldclim.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), with a spatial resolution of 5\u0026deg; longitude by 3.75\u0026deg; latitude (Dentener \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). We extracted AI from the Global Aridity Index and the Potential Evapotranspiration Database - Version 3, with a spatial resolution of 30 arc-seconds (Zomer et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). For edaphic factors, we extracted TOC, TN, TP, pH at 0\u0026ndash;30 cm soil depth from the global soil dataset for use in Earth system models, with a 1 km spatial resolution (Shangguan et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2 Statistical analysis\u003c/h2\u003e\u003cp\u003eIn this study, we selected the natural log-transformed response ratio (RR) to calculate the \u0026ldquo;effect size\u0026rdquo; of different treatments on hyphal and root traits (Hedges et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). We defined RR as the ratio of the value of a parameter in the treatment group to that in the control group. We employed a random effects model to calculate RR and to generate confidence intervals (CI) with the restricted maximum likelihood method using the \u003cem\u003erma.mv\u003c/em\u003e function in the metafor package version 4.8-0 (Viechtbauer \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). If the 95% CI did not overlap with zero, the effects of treatments on the response variables were considered statistically significant at \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05. We explored the possibility of publication bias using Egger\u0026rsquo;s regression test (\u003cem\u003eZ\u003c/em\u003e- and \u003cem\u003eP\u003c/em\u003e-values are given), trim and fill model and bootstrap approach (Egger et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Veroniki et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Rossetti et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) to examine the validity of the results of our meta-analysis. Moreover, RR and its CI were transformed back to the percentage change (% Change) for ease of interpretation. We employed linear regression and generalized additive models to evaluate the relationships between the effects of nitrogen/organic additions on roots and their effects on hyphae for each mycorrhizal type and for the aggregated dataset, respectively. Moreover, we used linear regression analysis to highlight the relationships of effect sizes of nitrogen/organic nutrient additions on roots and hyphae in relation to various soil and experimental variables.\u003c/p\u003e\u003cp\u003eMetaForest, a random forest-based algorithm, takes into account different weighting between experiments, incorporates several different predictors and their interactions, and considers the non-linear relationship between moderators and the predicted variables (van Lissa \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In this study, we used MetaForest analysis to determine the relative importance of climate, soil properties, mycorrhizal types, root sampling methods, and fertilization regimes in modulating the responses of roots and hyphae to nutrient additions. The analysis was conducted using the \u003cem\u003eMetaForest\u003c/em\u003e function in the MetaForest package version 0.1.4. Our analytical procedure initially involved assessing the convergence of the original model, followed by preselecting the moderators, tuning the model parameters, selecting the final model, and then verifying its convergence. We evaluated model performance using the retrodictive \u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003efit\u003c/em\u003e\u003c/sub\u003e) and the predictive \u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e metrics derived from cross-validation (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003ecv\u003c/em\u003e\u003c/sub\u003e) and out-of-bag testing (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003eoob\u003c/em\u003e\u003c/sub\u003e) (van Lissa \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e"},{"header":"3 Results","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003e3.1 Responses of hyphae and roots to nitrogen, phosphorus and organic nutrient additions\u003c/h2\u003e\u003cp\u003eNitrogen addition resulted in a significant increase in hyphal length (26%), root biomass (34%) and root length (31%), while concurrently causing a significant reduction in root colonization (-20%). Phosphorus addition led to notable enhancements in root biomass (27%) and root length (36%), but did not significantly influence hyphal biomass, hyphal length or root colonization. Organic nutrient addition significantly increased hyphal biomass (63%) and length (268%), as well as root biomass (87%) and root length (24%), without affecting root colonization (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Moreover, the combined application of nitrogen and phosphorus addition did not produce significant changes in any hyphal or root traits (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Notably, roots and hyphae associated with ectomycorrhizal hosts showed more pronounced responses to nutrient additions compared to those of arbuscular mycorrhizal hosts (Figure S1). Comprehensive analyses employing Egger\u0026rsquo;s regression test, the trim and fill method and bootstrap approaches confirmed that the directionality and statistical significance of effect sizes related to nitrogen, phosphorus, combined nitrogen and phosphorus, and organic nutrient addition remained robust despite potential publication bias (Table S2).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eLinear regression analyses revealed a positive linear relationship between the effects of nitrogen addition on root biomass and ectomycorrhizal hyphal length, whereas a negative linear relationship was observed between the effects of organic nutrient addition on root biomass and ectomycorrhizal hyphal length (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In contrast, no significant relationships were detected for arbuscular mycorrhizal hosts (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003e). When data were aggregated, the linear and nonlinear relationships emerged between the effects of nitrogen and organic nutrient additions on root biomass and their corresponding effects on hyphal length, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\u003ch2\u003e3.2 Key factors influencing hyphae and roots response to nitrogen and organic nutrient additions\u003c/h2\u003e\u003cp\u003eMetaForest modeling accounted for 45%, 71%, 83% and 51% of the variance in the effects of nitrogen addition on hyphal length, root biomass, root length and root colonization, respectively, identifying fertilization regime, climatic variables, and soil properties as important moderators (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Specifically, experiment duration and soil total phosphorus were the two most influential factors affecting the impact of nitrogen addition on hyphal length and root biomass (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, b), whereas soil total organic carbon and mean annual temperature, as well as amount of nitrogen addition and experiment duration were critical determinants for root length and root colonization, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e3\u003c/span\u003ec, d). The effects of nitrogen addition on hyphal length, root biomass and root colonization exhibited significant declines with increasing experiment duration/amount of nitrogen addition (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-c). Additionally, nitrogen-induced effects on root length and root colonization significantly decreased with mean annual temperature and latitude, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ed, f), while effects on root length significantly increased with latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). Effects of nitrogen addition on hyphal length, root biomass and root colonization were negatively correlated with soil total phosphorus or soil pH (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ea, b, d, h), whereas root length and hyphal length significantly increased with soil total phosphorus, soil total organic carbon and soil pH (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ec, e-g).\u003c/p\u003e\u003cp\u003eRegarding organic nutrient addition, MetaForest models explained 53% and 80% of the variation in their effects on organic nutrient addition on hyphal biomass and root biomass, respectively, with mycorrhizal type and soil properties (such as soil total nitrogen) serving as key moderators (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Figure S2a, b). Moreover, mycorrhizal type and soil total organic carbon emerged as the first influential factors governing the effects of phosphorus addition on root biomass and root length (Figure S2c, d).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003ePerformance of MetaForest models in predicting effects of nitrogen (N), phosphorus (P), N-P combination (NP) and organic nutrient (O) additions on hyphal biomass (HB), hyphal length (HL), root biomass (RB), root length (RL) and root colonization (RC). Model performance is evaluated using the retrodictive \u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003efit\u003c/em\u003e\u003c/sub\u003e), and the predictive \u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e values obtained during cross validation (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003ecv\u003c/em\u003e\u003c/sub\u003e) and out-of-bag tests (\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003eoob\u003c/em\u003e\u003c/sub\u003e). Non-negative \u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003eoob\u003c/em\u003e\u003c/sub\u003e values indicate no serious model overfitting.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"15\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003efit\u003c/em\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c10\" namest=\"c7\"\u003e\u003cp\u003e\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003ecv\u003c/em\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c15\" namest=\"c12\"\u003e\u003cp\u003e\u003cem\u003eR\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003csub\u003e\u003cem\u003eoob\u003c/em\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eNP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003eN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003eNP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003eO\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eHB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.53\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003e0.03\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eHL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.80\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.62\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003e0.59\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.83\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.51\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c15\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable S1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThe reference list (the details on references are provided in the data sources section at the conclusion of the text) regarding the classifications of terrestrial ecosystems, the types of mycorrhizae and the categories of nutrient additions utilized in this study.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eReferences\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eInorganic nutrient addition\u003c/span\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eOrganic nutrient addition (O)\u003c/span\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNitrogen (N)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePhosphorus (P) and wood ash (P)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eN-P combination (NP)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan type=\"BoldUnderline\" class=\"BoldUnderline\" name=\"Emphasis\"\u003eGrasses/Meadows, with AM\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYang Y, Zhang J, He J et al (2023)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCa(H\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e, Ca(H\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAntoninka A, Reich PB and Johnson NC (2011)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLiu Y, Shi G, Mao L et al (2012)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e(NH\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003eHPO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMuneer MA, Wang P, Zaib-un-Nisa et al (2020)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eCl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMuneer MA, Wang P, Zhang J et al (2020)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eCl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan type=\"BoldUnderline\" class=\"BoldUnderline\" name=\"Emphasis\"\u003eShrubs/Shrubland, with AM\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRillig MC and Allen MF (1998)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCaNO\u003csub\u003e3\u003c/sub\u003e, NH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePan S, Wang Y, Qiu Y et al (2020)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan type=\"BoldUnderline\" class=\"BoldUnderline\" name=\"Emphasis\"\u003eTrees/Crops/Forest, with AM\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYan G, Zhou M, Wang M et al (2019)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eQu Z, Zhu L, Jiang Q et al (2024)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u0026middot;2H\u003csub\u003e2\u003c/sub\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eChen W, Koide RT, Adams TS et al (2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCamenzind T, Homeier J, Dietrich K et al (2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u0026middot;2H\u003csub\u003e2\u003c/sub\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e, NaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u0026middot;2H\u003csub\u003e2\u003c/sub\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLiu B, Li L, Rengel Z et al (2019)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZheng C, Chai M, Jiang S et al (2015)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eKH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLi L, McCormack ML, Chen F et al (2019)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e, NaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCheng L, Chen W, Adams TS et al (2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e, NaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBethlenfalvay GJ (1983)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCa\u003csub\u003e10\u003c/sub\u003e(PO\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e6\u003c/sub\u003e(OH)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFrey JE and Ellis JR (1997)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, NH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStaddon PL, Jakobsen I and Blum H (2004)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eHodge A and Fitter AH (2010)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCao C (2021)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZhang F (2019)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eChen W, Koide RT and Eissenstat DM (2018)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan type=\"BoldUnderline\" class=\"BoldUnderline\" name=\"Emphasis\"\u003eTrees/Forest, with ECM\u003c/span\u003e:\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAlberton O and Kuyper TW (2009)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYan G, Zhou M, Wang M et al (2019)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZhang J, Lin G and Zeng D-H (2024)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eChen W, Koide RT, Adams TS et al (2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEkblad A, Wallander H, Carlsson R et al (1995)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eKH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, K\u003csub\u003e2\u003c/sub\u003eHPO\u003csub\u003e4\u003c/sub\u003e, NaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, Na\u003csub\u003e2\u003c/sub\u003eHPO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, KH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, K\u003csub\u003e2\u003c/sub\u003eHPO\u003csub\u003e4\u003c/sub\u003e, NaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e, Na\u003csub\u003e2\u003c/sub\u003eHPO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eK\u0026aring;r\u0026eacute;n O and Nylund JE (1996)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e(NH\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLeppalammi-Kujansuu J, Ostonen I, Stromgren M et al (2013)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWeigt RB, Raidl S, Verma R et al (2011)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWiemken V, Ineichen K and Boller T (2001)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCheng L, Chen W, Adams TS et al (2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e, KH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMajdi H, Truus L, Johansson U et al (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2008\u003c/span\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eWood ash granules\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eJones MD, Durall DM and Tinker PB (1990)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eKH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePampolina NM, Dell B and Malajczuk N (2002)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCa(H₂PO₄)\u003csub\u003e2\u003c/sub\u003e\u0026middot;CaHPO₄\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTreseder KK, Turner KM and Mack MC (2007)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBakker MR, Jolicoeur E, Trichet P et al (2009)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNaples BK and Fisk MC (2010)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaH\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRasheed MU, Kasurinen A, Kivim\u0026auml;enp\u0026auml;\u0026auml; M et al (2017)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eYara Peatcare TM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZhu X, Zhang Z, Wang Q et al (2022)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWang C, Brunner I, Guo W et al (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eHe Y (2021)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCa(H\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u0026middot;H\u003csub\u003e2\u003c/sub\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWang G (\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2021\u003c/span\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eOsmocote\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChicken manure\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eForsmark B, Nordin A, Rosenstock NP et al (2021)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDu Z, Wang W, Zeng W et al (2014)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCO(NH\u003csub\u003e2\u003c/sub\u003e)\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eChen W, Koide RT and Eissenstat DM (2018)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLeaf\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable S2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eResults of testing publication bias for statistically significant response variables for nitrogen (N), phosphorus (P), N-P combination (NP) and organic nutrient (O) additions on hyphal biomass (HB), hyphal length (HL), root biomass (RB), root length (RL) and root colonization (RC) using Egger\u0026rsquo;s regression test (\u003cem\u003eZ\u003c/em\u003e- and \u003cem\u003eP\u003c/em\u003e-values are given), trim and fill model and bootstrap approach (5% and 95% confidence intervals [CI] of effect sizes are given).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"13\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eEgger\u0026rsquo;s regression\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003eTrim and fill model\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003eBootstrap approach\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e\u003cp\u003eOriginal model\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTreatments\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVariables\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003e\u003cem\u003eZ P\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e5%CI 95%CI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e5%CI 95%CI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e\u003cp\u003e5%CI 95%CI\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-1.213\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.225\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.063\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.252\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.118\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.112\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.149\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.148\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.486\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.137\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.152\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.403\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.181\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.376\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.152\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.403\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-1.874\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.060\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.443\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.770\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.174\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.417\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.457\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.011\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.010\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.293\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.391\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.122\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.413\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-3.261\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.264\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-0.025\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.302\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e-0.137\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.329\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e-0.120\u003c/p\u003e \u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3.019\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.221\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.152\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.166\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.111\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.221\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.152\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.180\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.238\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.034\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.307\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.011\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.271\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.034\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.307\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.175\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.861\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.161\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.419\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.136\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.357\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.113\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.371\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.440\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.660\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.317\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.724\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.175\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.461\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.123\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.489\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-2.623\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.009\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.358\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.014\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.324\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e-0.020\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.358\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.014\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eNP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.937\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.349\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.287\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" 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colname=\"c3\"\u003e\u003cp\u003e0.974\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.330\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.283\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.054\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.117\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.114\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.158\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.154\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eO\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.374\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.196\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.684\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.214\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.689\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" 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colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e1.244\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.361\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-1.713\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.087\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.193\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.320\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.952\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.034\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.865\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.065\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.267\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.102\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.333\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.068\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.367\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.342\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.732\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.051\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.142\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.050\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.137\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e-0.056\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.138\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"4 Discussion","content":"\u003cp\u003e\u003cb\u003e4.1 Variations in nutrient acquisition of roots and hyphae in response to inorganic and organic nutrient additions\u003c/b\u003e\u003c/p\u003e\u003cp\u003eThis study revealed that roots exhibited more pronounced positive responses to the addition of inorganic nutrients, specifically nitrogen and phosphorus compared to hyphae (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e), supporting our first hypothesis. Typically, plants tend to prioritize root proliferation under conditions of abundant inorganic nutrient availability, thereby employing an inorganic nutrient acquisition strategy (Mohan et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Chari et al. 2024; Wang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Although hyphal length significantly increased following nitrogen addition, a concurrent decrease in root colonization was observed, suggesting that plants may reduce their reliance on mycorrhizal associations while enhancing direct nitrogen uptake through roots under elevated nitrogen inputs (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e6\u003c/span\u003e). However, in comparison to roots, mycorrhizal fungi require a greater carbon investment due to their rapid hyphal turnover rates (Jakobsen and Rosendahl \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Consequently, plants reduce the carbon maintenance costs associated with nitrogen acquisition by allocating more carbon to fine roots and less to mycorrhizal hyphae as soil nitrogen availability increases, particularly in regions characterized by lower temperatures and higher latitudes (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ed-f). Furthermore, phosphorus addition significantly increased root length and root biomass but did not affect hyphal length, hyphal biomass or root colonization (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Previous studies have demonstrated that plants can alter their root morphology to optimize inorganic phosphorus acquisition by producing more finely branched roots and elongating root hairs (Lambers et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; White et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; P\u0026eacute;ret et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These results suggest that plants with an adequate supply of inorganic phosphorus prefer to use their roots for phosphorus uptake, as they exhibit higher uptake efficiency and physiological performance compared to phosphorus-deficient conditions, where mycorrhizal symbiosis plays a more important role (Brown et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Consequently, plants employ root-dependent nutrient strategies in response to elevated inorganic nitrogen and phosphorus availability (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn this study, we observed an increase in root biomass and root length after the addition of organic nutrients, suggesting that plants may release root exudates that stimulate microbial priming effects, thereby promoting the weathering of soil minerals and the mineralization of soil organic matter, and thus also the release of the associated nutrients (Li et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Nonetheless, the overall root responses to organic addition were relatively modest compared to the pronounced positive effects observed on fungal hyphae (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e), again confirming our first hypothesis. The enhancement of fungal hyphae due to organic nutrient addition is attributed to the beneficial impacts of increased organic matter on soil water status, soil structure and synergistic microbial activity, and the reduction of mechanical resistance to hyphal penetration within the soil (Ryan et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Joner and Jakobsen \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Gryndler et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Hammer et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). As expected in our second hypothesis, our findings indicate that soil total organic carbon is a critical factor modulating the effects of nitrogen addition on both hyphal length and root length, with positive correlations observed between these variables and soil total organic carbon (Figs.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, c and \u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ee, f). Soil organic carbon contributes to the stabilization of soil structure and enhances soil cation exchange capacity, both of which are essential for soil nutrient retention within the soil (Chen et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Consequently, soils with a higher soil organic carbon content are able to retain larger amounts of plant-available nitrogen under the same nitrogen additions, thereby eliciting a more pronounced root response to the nitrogen supplementation. Moreover, the accelerated growth observed in soils enriched with the higher organic matter may confer a selective advantage to fungal hyphae in forest ecosystems frequently subjected to disturbance such as wildfires, facilitating enhanced nutrient acquisition from post-fire ash deposits (Majdi et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The decomposition of organic matter in close proximity to the mycorrhizal hyphae provides a rich source of accessible carbon and energy (Gryndler et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Accordingly, plants may preferentially adopt an organic nutrient acquisition strategy, wherein symbiotic associations promote microbial mineralization of nitrogen and phosphorus under conditions characterized by abundant organic nutrient availability (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003e4.2 Factors influencing root and hyphal responses to inorganic and organic nutrient additions\u003c/h2\u003e\u003cp\u003eIn this meta-analysis, a positive correlation was identified between the effects of nitrogen addition on root biomass and those on hyphal length in ectomycorrhizal hosts (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). This associations is consistent with the premise that root and hyphal development are interconnected, as hyphal growth is driven by carbon allocation from fine roots, thereby fostering a positive relationship between root formation and fungal hyphae proliferation (Korkama et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Elevated nitrogen deposition mitigates nitrogen limitation within ecosystems, thereby enhancing plant productivity, which is indicative of a greater allocation of carbon belowground, specifically to roots and their symbiotic mycorrhizae (Schulte-Uebbing and De Vries \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Nonetheless, plants must strategically allocate carbon resources between root-mediated nutrient uptake and hyphal nutrient acquisition mechanisms under nutrient-limited conditions (Wang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). A prominent illustration of this resource allocation trade-off is the antagonistic interaction observed between mycorrhizal symbioses and fine-root growth, attributable to their morphological similarities and potential functional redundancy (Brown et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Bergmann et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This antagonism is corroborated by the significantly negative correlation found between the effects of organic nutrient amendments on root biomass and hyphal length in ectomycorrhizal hosts (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). Compared to roots, ectomycorrhizal fungi exhibit greater efficiency in extracting nutrients from soil organic matter (Brown et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Collectively, these findings suggest that ectomycorrhizal plants may intensify their symbiotic interactions with mycorrhizal fungi, relying less on the proliferation of fine roots to optimize the acquisition of organic nutrients following organic nutrient amendments.\u003c/p\u003e\u003cp\u003eIn the present study, no significant correlation was observed between the effects of organic nutrient amendments on root biomass and their effects on hyphal length in arbuscular mycorrhizal hosts (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). These results suggest that the influence of organic nutrient additions on root and hyphae development depends on the type of mycorrhiza (Figures S1, S2). The divergent nutrient acquisition strategies exhibited by arbuscular mycorrhizal and ectomycorrhizal hosts in response to organic nutrient amendments may be attributed to variations in environmental conditions and/or differential capacities to access nutrients from soil organic matter (Chari et al. 2024; Guo et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Unlike arbuscular mycorrhizal hosts, which are predominantly distributed across tropical and subtropical regions, ectomycorrhizal hosts primarily inhabit temperate and boreal zones characterized by soils rich in organic matter but typically deficient in inorganic nitrogen (Reich and Oleksyn \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Consequently, ectomycorrhizal fungi generally exhibit superior capabilities in mineralizing soil organic matter compared to arbuscular mycorrhizal fungi (Rosling et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The findings of this study underscore that the distinct responses of roots and hyphae to organic nutrient amendments, differentiated by mycorrhizal types, can substantially affect resource acquisition. Nonetheless, further research is required to elucidate the underlying mechanisms driving these observed patterns on arbuscular mycorrhizal \u003cem\u003eversus\u003c/em\u003e ectomycorrhizal host plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe MetaForest analysis identified fertilization regime as the primary determinant influencing the impact of nitrogen addition on root and hyphal growth (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e3\u003c/span\u003e), being supported by our second hypothesis. Specifically, short-term nitrogen supplementation appears to promote the development of roots and hyphae; however, this beneficial effect diminishes with extended nitrogen application, particularly beyond a decade. This pattern can be elucidated by several mechanisms. Initially, nitrogen addition enhances plant growth and increases carbon allocation to belowground structures, but these effects decline as the duration of nitrogen exposure lengthens (Li et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Song et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). This observation aligns with findings from a recent meta-analysis demonstrating a temporal decrease in nitrogen-induced fungal biomass enhancement (Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Furthermore, prolonged nitrogen addition may lead to soil acidification, which adversely affects root and microbial proliferation, thereby counteracting the initial positive effects of nitrogen enrichment (Sterkenburg et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This is corroborated by the significant positive correlation observed between nitrogen addition effects on root length and soil pH (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003eg). Moreover, our observations indicate that root colonization markedly declines as the amount of nitrogen supplementation increases (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ec), implying that plants predominantly acquire nutrients through fine roots rather than mycorrhizal fungi when nitrogen availability in the soil is elevated (Ma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Taken together, these convergent lines of evidence underscore fertilization regime, such as experiment duration and amount of nutrient addition, as a critical factor modulating both the direction and magnitude of nitrogen addition effects on roots and hyphae.\u003c/p\u003e\u003cp\u003eThis study identified soil total phosphorus as a key factor influencing the variability in hyphal and root responses to nitrogen amendment (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-d). Nitrogen addition promoted hyphal proliferation, as evidenced by increased hyphal length and root colonization under conditions of low phosphorus availability (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ea, d). Generally, the abundance of root-external hyphae is positively associated with phosphorus acquisition efficiency in mycorrhizal plants (de Miranda and Harris \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Sbrana et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Consequently, the initial enhancement of hyphal length and root colonization partially mitigates phosphorus limitation for plant growth as soil nitrogen availability rises. Negative growth responses in mycorrhizal hyphae may occur when carbon-limited fungi must allocate a greater proportion of host-derived carbon to excessive phosphorus assimilation rather than to growth. Conversely, nitrogen addition exerts more pronounced positive effects on root growth, as indicated by increased root length in areas with higher phosphorus concentrations (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ec). Collectively, these results suggest a shift in plant nutrient acquisition strategies from reliance on mycorrhizal associations toward greater dependence on fine roots (e.g., increased root length observed in this study) with increasing soil phosphorus levels. Thus, plants appear to strategically allocate limited carbon resources between root and mycorrhizal traits, balancing these two principal pathways for phosphorus uptake (Wang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). However, this proposed mechanism could not be confirmed through combined nitrogen and phosphorus additions due to insufficient paired data in the present study. Therefore, future research should prioritize investigating plant mycorrhizal symbioses and fine-root traits in response to simultaneous nitrogen and phosphorus supplementation.\u003c/p\u003e\u003c/div\u003e"},{"header":"5. Conclusions","content":"\u003cp\u003eOur meta-analysis elucidated the complex responses of roots and fungal hyphae to nitrogen, phosphorus, and organic nutrient additions, indicating shifts in plant nutrient acquisition strategies in relation to the availability of inorganic and/or organic soil nutrients. The results demonstrated that roots possess advantages over hyphae in terms of root length and biomass, thereby exhibiting greater adaptability to inorganic nutrient (nitrogen and phosphorus) supplementation. Conversely, fungal hyphae, particularly those associated with ectomycorrhizal hosts, showed enhanced growth and distinct advantages relative to roots following organic nutrient additions. Additionally, a significant positive correlation was observed between the effect sizes on root biomass and ectomycorrhizal hyphal length under nitrogen addition, whereas a negative correlation was found under organic addition. Moreover, our findings suggest that fertilizer regimes (e.g. experiment duration and amount of nutrient addition), climatic factors (e.g. mean annual temperature), soil characteristics (e.g. pH, phosphorus, and organic carbon concentrations), and mycorrhizal types collectively influence the responses of roots and hyphae to nitrogen and organic amendments. These divergent responses underscore the necessity of integrating both root and hyphal dynamics within nutrient acquisition frameworks to accurately evaluate and predict the consequences of global environmental changes on carbon, nitrogen, and phosphorus cycling.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003ch2\u003eDeclaration of interests\u003c/h2\u003e\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e\u003cp\u003eThis work was financially supported by the Natural Science Foundation of China (grant number 42171051).\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eBergmann J, Weigelt A, van der Plas F et al (2020) The fungal collaboration gradient dominates the root economics space in plants. 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Pak J Bot 51:727\u0026ndash;733. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.30848/PJB2019-2(39)\u003c/span\u003e\u003cspan address=\"10.30848/PJB2019-2(39)\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"mycorrhizal, nitrogen, nutrient acquisition, phosphorus, soil organic matter","lastPublishedDoi":"10.21203/rs.3.rs-7788604/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7788604/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground and Aims\u003c/h2\u003e\u003cp\u003eFine roots and mycorrhizal hyphae represent two critical pathways for plants to acquire nutrients. Nevertheless, the mechanisms by which roots and hyphae respond to variations in inorganic and organic nutrients remain inadequately understood.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e\u003cp\u003eWe conducted a meta-analysis on the effects of nitrogen, phosphorus, nitrogen-phosphorus combination and organic nutrient additions on root length (RL), root biomass (RB), root colonization (RC), hyphal length (HL), and hyphal biomass (HB) based on 738 root-hyphae observation pairs.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e\u003cp\u003eWe found that RB and RL increased by 27%-34% and 31%-36%, respectively, following nitrogen and/or phosphorus additions, whereas HL increased by 26% and RC decreased by 20%. The effect sizes of organic nutrient addition on hyphae were two to three times greater than those observed for fine roots. The impact of nitrogen addition on HL, RB and RC significantly decreased with increasing experiment duration, amount of nitrogen addition and/or soil total phosphorus, conversely, its effect on HL and RL significantly intensified with soil total organic carbon. The effects of nitrogen and organic nutrient additions on RB exhibited significant positive and negative correlations, respectively, with their effects on HL in ectomycorrhizal hosts. No significant relationships were detected for arbuscular mycorrhizal hosts.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e\u003cp\u003eOur results demonstrate that plants preferentially employ root-dependent acquisition strategies under high inorganic nutrients but rely more on hypha-dependent strategies under abundant organic nutrients. This study underscores the necessity of integrating both roots and hyphae within current nutrient acquisition frameworks to evaluate the effects of global changes on carbon and nitrogen cycling.\u003c/p\u003e","manuscriptTitle":"Asymmetric responses of roots and hyphae to inorganic and organic nutrient additions: a global meta-analysis","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-22 21:36:47","doi":"10.21203/rs.3.rs-7788604/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2025-11-29T03:38:15+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-10-15T07:30:36+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-09T02:26:09+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant and Soil","date":"2025-10-09T01:18:17+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-10-08T23:56:41+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant and Soil","date":"2025-10-06T02:37:32+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"bb0e1d24-041b-47d2-bc4c-32aaca788135","owner":[],"postedDate":"October 22nd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-23T16:09:53+00:00","versionOfRecord":{"articleIdentity":"rs-7788604","link":"https://doi.org/10.1007/s11104-026-08389-8","journal":{"identity":"plant-and-soil","isVorOnly":false,"title":"Plant and Soil"},"publishedOn":"2026-02-20 15:59:19","publishedOnDateReadable":"February 20th, 2026"},"versionCreatedAt":"2025-10-22 21:36:47","video":"","vorDoi":"10.1007/s11104-026-08389-8","vorDoiUrl":"https://doi.org/10.1007/s11104-026-08389-8","workflowStages":[]},"version":"v1","identity":"rs-7788604","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7788604","identity":"rs-7788604","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}