Global pattern and controls of biological nitrogen fixation under nutrient enrichment: A meta‐analysis

Biological nitrogen (N) fixation (BNF), an important source of N in terrestrial ecosystems, plays a critical role in terrestrial nutrient cycling and net primary productivity. Currently, large uncertainty exists regarding how nutrient availability regulates terrestrial BNF and the drivers responsible for this process. We conducted a global meta‐analysis of terrestrial BNF in response to N, phosphorus (P), and micronutrient (Micro) addition across different biomes (i.e, tropical/subtropical forest, savanna, temperate forest, grassland, boreal forest, and tundra) and explored whether the BNF responses were affected by fertilization regimes (nutrient‐addition rates, duration, and total load) and environmental factors (mean annual temperature [MAT], mean annual precipitation [MAP], and N deposition). The results showed that N addition inhibited terrestrial BNF (by 19.0% (95% confidence interval [CI]: 17.7%?20.3%); hereafter), Micro addition stimulated terrestrial BNF (30.4% [25.7%?35.3%]), and P addition had an inconsistent effect on terrestrial BNF, i.e., inhibiting free‐living N fixation (7.5% [4.4%?10.6%]) and stimulating symbiotic N fixation (85.5% [25.8%?158.7%]). Furthermore, the response ratios (i.e., effect sizes) of BNF to nutrient addition were smaller in low‐latitude (<30°) biomes (8.5%?36.9%) than in mid‐/high‐latitude (≥30°) biomes (32.9%?61.3%), and the sensitivity (defined as the absolute value of response ratios) of BNF to nutrients in mid‐/high‐latitude biomes decreased with decreasing latitude (p ≤ 0.009; linear/logarithmic regression models). Fertilization regimes did not affect this phenomenon (p > 0.05), but environmental factors did affect it (p < 0.001) because MAT, MAP, and N deposition accounted for 5%?14%, 10%?32%, and 7%?18% of the variance in the BNF response ratios in cold (MAT < 15°C), low‐rainfall (MAP < 2,500 mm), and low‐N‐deposition (<7 kg ha^?1 year^?1) biomes, respectively. Overall, our meta‐analysis depicts a global pattern of nutrient impacts on terrestrial BNF and indicates that certain types of global change (i.e., warming, elevated precipitation and N deposition) may reduce the sensitivity of BNF in response to nutrient enrichment in mid‐/high‐latitude biomes.     

The Inhibitory Effects of Nitrogen Deposition on Asymbiotic Nitrogen Fixation are Divergent Between a Tropical and a Temperate Forest

Ecosystems - Asymbiotic nitrogen (N) fixation (ANF) is an important source of N in pristine forests and is predicted to decrease with N deposition. Previous studies revealing N fixation in response...     

Substrate stoichiometry determines nitrogen fixation throughout succession in southern Chinese forests

The traditional view holds that biological nitrogen (N) fixation often peaks in early‐ or mid‐successional ecosystems and declines throughout succession based on the hypothesis that soil N richness and/or phosphorus (P) depletion become disadvantageous to N fixers. This view, however, fails to support the observation that N fixers can remain active in many old‐growth forests despite the presence of N‐rich and/or P‐limiting soils. Here, we found unexpected increases in N fixation rates in the soil, forest floor, and moss throughout three successional forests and along six age‐gradient forests in southern China. We further found that the variation in N fixation was controlled by substrate carbon(C) : N and C : (N : P) stoichiometry rather than by substrate N or P. Our findings highlight the utility of ecological stoichiometry in illuminating the mechanisms that couple forest succession and N cycling.     

矿质养分输入对森林生物固氮的影响

生物固氮是森林生态系统重要的氮素来源,并且在全球氮循环中占有重要的地位。近代以来,因人类活动加剧而导致氮沉降的增加以及其它矿质养分元素(如磷、钼、铁等)输入的改变已成为影响森林生态系统生物固氮的重要因素之一,并引起了学术界的普遍关注。综述了国内外关于森林生物固氮对矿质养分输入的响应及机理。主要内容包括:(1)森林生物固氮的概念及主要的测定方法;(2)矿质养分输入对森林生物固氮的影响。整体上讲,氮素输入抑制了森林生物固氮,磷和其他营养元素输入则表现为促进作用。氮和磷、磷和微量元素同时添加均提高了森林的固氮量;(3)矿质养分改变森林生物固氮的机理。包括生物作用机制(如改变地表层固氮菌的数量或群落丰度、改变结瘤植物的根瘤生物量和附生植物的丰度或盖度)和环境作用机制(如引起土壤酸化、改变碳源物质的含量);(4)探讨了矿质养分输入对森林生物固氮影响研究中所存在的问题,并对未来该领域的研究提出建议。     

Effects of phosphorus addition with and without nitrogen addition on biological nitrogen fixation in tropical legume and non-legume tree plantations

It is well documented that phosphorus (P) input stimulates biological nitrogen (N) fixation (BNF) in tropical forests with non-legume trees. However, in tropical legume forests with soil N enrichment and P deficiency, the effects of P availability and its combination with N on BNF remain poorly understood. In this study, we measured BNF rate in different compartments, i.e., bulk soil, forest floor, rhizosphere, and nodules, in two tropical plantations with legume trees Acacia auriculiformis (AA) versus non-legume trees Eucalyptus urophylla, (EU) in southern China after 4 years of P addition and combined N and P additions. The objective was to investigate how P addition and its combination with N addition regulate BNF in a tropical legume plantation, and to compare the effects with those in a non-legume plantation. Our results showed that total BNF rates were significantly higher in the P-addition plots than in the control plots by 27.4 ± 4.3 and 23.3 ± 1.7 % in the EU and AA plantations, respectively. Total BNF rates were significantly higher in the NP-addition plots than in the control plots by 27.7  ± 5.0 and 8.5 ± 1.4 % in the EU and AA plantations, respectively, which contrasted to our previous result that total BNF rates were significantly lower in N-addition plots than in the control plots in the AA plantation. These findings suggest that P input can stimulate BNF in tropical forest biome dominated by legume trees, even in consideration of elevated atmospheric N deposition. Thus, our study revealed the important role of P in regulating biological N input, which should be taken into account in the modeling of biogeochemical cycles in the future.     

Effects of long-term nitrogen deposition on phosphorus leaching dynamics in a mature tropical forest

Elevated anthropogenic nitrogen (N) deposition is suggested to affect ecosystem phosphorus (P) cycling through altered biotic P demand and soil acidification. To date, however, there has been little information on how long-term N deposition regulates P fluxes in tropical forests, where P is often depleted. To address this question, we conducted a long-term N addition experiment in a mature tropical forest in southern China, using the following N treatments: 0, 50, 100, and 150 kg N ha^?1 year^?1. We hypothesized that (i) tropical forest ecosystems have conservative P cycling with low P output, and (ii) long-term N addition decreases total dissolved phosphorus (TDP) leaching losses due to reduced litter decomposition rates and stimulated P sorption deriving from accelerated soil acidification. As hypothesized, we demonstrated a closed P cycling with low leaching outputs in our forest. Under experimental N addition, TDP flux in throughfall was significantly reduced, suggesting that N addition may result in a less internal P recycling. Contrary to our hypothesis, N addition did not decrease TDP leaching, despite reduced litter decomposition and accelerated soil acidification. We find that N addition might have negative impacts on biological P uptake without affecting TDP leaching, and that the amount of TDP leaching from soil could be lower than a minimum concentration for TDP retention. Overall, we conclude that long-term N deposition does not necessarily decrease P effluxes from tropical forest ecosystems with conservative P cycling.     

氮、磷添加对不同林型土壤磷酸酶活性的影响

研究了鼎湖山3种森林类型(南亚热带季风常绿阔叶林、马尾松人工林和针叶阔叶混交林)的土壤酸性磷酸单酯酶活性(APA)对施肥的响应情况。在3种林型中分别设置对照、加氮(150 kg N hm-2a-1)、加磷(150 kg P hm-2a-1)以及N和P同时添加(150 kg N hm-2a-1+150 kg P hm-2a-1)4种不同处理。结果表明,季风林土壤APA((15.83±2.46)μmol g-1h-1)显著高于混交林((10.71±0.78)μmol g-1h-1)和马尾松林((9.12±0.38)μmol g-1h-1),且3种林型土壤APA与土壤有效磷含量均呈显著负相关。施加N肥显著提高了季风林土壤APA,而对混交林和马尾松林的作用不显著。施加P肥显著降低了混交林和马尾松林土壤APA,但对季风林的影响不明显。N和P同时添加仅显著降低了马尾松林土壤APA,但在季风林中存在交互作用。因此,N沉降会加剧亚热带成熟林土壤P的限制,可以考虑施加P肥作为森林管理的一种方式来缓解这种限制作用。