陆地生态系统植物多样性对矿质元素输入的响应

人类活动的加剧改变了陆地生态系统矿质元素(如氮、磷、钾等)循环的速度和方向,并且对生态系统的结构和功能也产生重要影响。如今,矿质元素输入量的改变及其产生的后续效应对陆地生态系统生物多样性的影响备受学者们的关注。从4个方面综述了全球氮沉降背景下主要矿质元素输入的改变对陆地植物多样性的影响及其机理:1)矿质营养元素限制的概念、确定方法以及与植物多样性的耦合关系;2)概述了氮、磷、钾等主要矿质元素输入对陆地植物多样性的影响:主要表现为负面效应;3)探讨了矿质元素输入影响植物多样性的可能机制,包括生态系统水平上的机制(如竞争排斥、酸化铝毒、物种入侵、同质性假说,间接诱导机制等)和植物个体水平上的机制(如元素失衡和环境敏感性增加等);4)根据目前研究现状,指出了已有研究的局限性,分析了未来可能的研究方向和重点。     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响, 尤其是加剧了植物生长的磷限制, 从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响, 认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量, 有利于植物的磷吸收并增加植物磷含量。同时, 森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性, 该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制: 1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力; 3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述, 外源氮输入使植物磷含量发生改变, 首要原因是土壤有效磷含量的改变, 其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响,尤其是加剧了植物生长的磷限制,从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响,认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量,有利于植物的磷吸收并增加植物磷含量。同时,森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性,该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制:1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力;3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述,外源氮输入使植物磷含量发生改变,首要原因是土壤有效磷含量的改变,其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

氮磷输入对过度放牧退化草原土壤线虫群落的影响

线虫是陆地生态系统中数量最多的一类无脊椎后生动物,在土壤碎屑食物网中占据多个营养级,并在能量流动和养分循环中发挥重要的生态功能.土壤线虫的群落结构和多样性水平可以反映生态系统功能的变化.多年来,过度放牧等不合理的开发利用导致我国大面积的草原处在不同程度的退化之中,外源性养分输入是维持其养分平衡进而恢复其生态和生产功能的重要措施.本研究以内蒙古锡林郭勒退化草原为对象,就氮磷输入对土壤线虫丰度和群落结构的单独效应和交互效应进行了研究.本研究获得线虫38属,其中垫咽属、滑刃属、索努斯属和盾垫属是该退化草原线虫群落优势属.氮输入降低了土壤线虫总丰度、捕食杂食线虫丰度以及植物寄生线虫丰度,而磷输入提高了土壤线虫总丰度、食真菌线虫丰度、捕食杂食类线虫丰度以及植物寄生线虫丰度,氮输入抑制了磷输入对线虫总丰度、捕食杂食线虫和植物寄生线虫丰度的促进作用.氮磷输入对线虫多样性无影响,可能与不同养分输入下稳定的植物群落多样性有关.氮输入显著提升了退化草原线虫成熟度指数,降低了植物寄生线虫成熟度指数(PPI),并且能缓解磷输入对PPI和瓦斯乐斯卡指数的负面作用,表明氮输入提高退化草地土壤健康程度,促进线虫群落和食物网的稳定性.上述研究结果有助于从土壤生物学角度理解氮磷输入对退化草地恢复的影响机理.     

氮沉降对森林土壤磷循环的影响

磷是生物体必需的大量元素之一,也是许多生态系统的主要限制因子。近年来,大气氮沉降日益加剧,已对森林生态系统磷循环产生了不可忽视的影响。关于氮沉降对生态系统磷循环的影响已开展了一系列的研究,然而尚缺少对其整体的认识。因此,通过收集国内外已发表的相关文章,从以下五个方面综述氮沉降对森林生态系统土壤磷循环的影响及其机理:1)阐述了森林生态系统土壤磷循环的概念;2)介绍了氮沉降对森林土壤磷循环影响的研究方法,包括长期定位模拟氮沉降法、自然氮沉降梯度法和同位素示踪法等;3)概述了氮沉降对森林生态系统土壤磷循环的影响。目前的研究结论趋向于认为长期氮沉降使森林土壤磷循环速率加快。长期氮输入易于使土壤中可溶性磷向非活性磷酸盐库迁移而难以被利用。因此,为了满足需求,土壤磷酸酶活性将增加以加速有机磷的矿化,从而加速磷素在土壤-植物-微生物之间的周转。4)探讨了氮沉降影响森林土壤磷循环的机制。氮沉降可通过改变土壤有机质的性质、微生物群落组成、磷酸酶活性以及阳离子的流动性等途径影响森林土壤磷循环;5)指出了当前研究存在的问题及未来的研究方向。     

林火干扰对森林生态系统土壤有机碳的影响研究进展

林火干扰是森林生态系统特殊而重要的生态因子,可改变生态系统的养分循环与能量传递。研究林火干扰对森林生态系统土壤有机碳的影响,有助于理解森林生态系统中土壤碳固持和碳循环过程,为制定科学合理的旨在减缓全球变化的林火管理策略具有重要意义。从4个方面阐述了林火干扰对森林生态系统土壤有机碳的影响及内在机制:分别从大尺度和小尺度两个方面阐述了林火干扰对土壤有机碳的影响及对森林生态系统碳循环与碳平衡的作用机制;探讨了不同林火干扰类型和林火干扰强度下,土壤活性有机碳对林火干扰的响应机制;阐明了林火干扰对土壤惰性有机碳的影响及作用机制;论述了林火干扰主要通过改变土壤有机碳的输入和输出过程进而影响土壤有机碳的稳定性及内在机制。最后提出了提高林火干扰对森林生态系统土壤有机碳影响定量化研究的4种路径选择:(1)全面比较研究不同林火干扰类型对土壤有机碳循环及其碳素再分配过程的功能特征;(2)进一步阐明林火干扰通过改变植被结构进而影响土壤生物群落结构,剖析土壤碳库循环的内在机制;(3)完善不同时空尺度下林火干扰对森林生态系统土壤碳库周转过程的定量化研究;(4)加强不同林火干扰类型土壤碳库稳定性差异的研究。     

氮磷添加对树木生长和森林生产力影响的研究进展

人为活动所导致的氮、磷输入和大气氮、磷沉降使生态系统中的氮、磷可利用性大幅提高, 对陆地生态系统的碳循环过程产生了显著影响。树木生长和森林生产力在全球碳循环中发挥着重要作用, 它决定着陆地碳固存的大小和方向。目前, 在全球范围内开展了很多氮、磷添加调控树木生长和森林生产力的野外控制实验, 但是研究结果并不一致, 受到多种生物、环境和实验处理条件等因素的影响。该文从野外氮添加和磷添加实验的文献数量、实验数量及其全球空间分布三个方面概述了氮、磷添加对树木生长和森林生产力影响的研究现状, 并总结了氮、磷添加实验中树木生长和森林生产力的评估方法, 包括相对生长速率和绝对增长量。基于相关的研究结果, 阐述了氮、磷添加影响树木生长和森林生产力的调控因素及其潜在影响机制, 包括气候、树木径级与林龄、植物功能性状(共生菌根类型、树木固氮属性和保守性与获得性性状)、植物和微生物相互作用关系、区域养分沉降速率和实验处理条件等。最后, 基于当前的研究进行了系统总结, 并指出今后需要加强的几个方面的研究, 以期为后续研究提供参考: 树木生长响应氮、磷添加的生理学机制, 树木各部分生长对氮、磷添加响应的权衡与分配, 植物功能性状在调节与预测树木生长响应氮、磷添加中的作用, 树木之间的竞争关系如何调控氮、磷添加对树木生长的影响, 以及开展长期的和联网的氮、磷添加对树木生长和森林生产力影响的野外控制实验。     

Nitrogen and phosphorus translocation of forest floor mosses as affected by a pulse of these nutrients

藓类是亚高山森林、北方针叶林和极地苔原等原始生态系统的重要组成部分。养分沉降是这些生态系统的主要养分来源。然而,对 藓类如何利用大气沉降这一时间上异质性较高的养分来源的认识缺乏,限制了我们对藓类在生态系统养分循环中作用的认识,也制约了模 拟和预测生态系统服务功能动态变化的有效性。本研究中,我们重点关注如下问题：(1)藓类新生部分的氮、磷含量随氮或磷沉降量的变化; (2)氮或磷养分沉降如何影响藓类的氮磷转运;(3)氮磷转运与藓类新生部分养分含量的关联性？为此,我们在位于青藏高原东部的亚高山 冷杉林中开展了设置有不同氮、磷施加量水平的突发养分沉降实验。以林下优势度最高的塔藓和锦丝藓为对象,我们在施加实验处理8天和1年后,分别测定了不同年龄藓类茎段中的氮、磷含量。研究结果发现,养分处理8天后,锦丝藓和塔藓的所有茎段的氮或磷含量均随处理中氮或磷施加量的增加而增加,表明它们能够高效率地吸收施加的氮或磷。养分处理1年后,形成于养分处理后的新生茎段中的氮、磷含量以及氮磷比与处理后8天的藓类茎段中氮、磷含量以及氮磷比呈正相关关系,表明藓类将吸收的养分转运至新生部分,转运过程受源—库关系的调控。磷转运效率解释了新生茎段21%–23%的氮磷比变化,而氮转运与新生茎段氮磷比没有显著关系,结果暗示磷转运对藓类生长的重要性。上述结果表明,养分转运是藓类植物利用间断的沉降养分的关键机制,也使藓类成为生态系统中重要的养分库。     

氮输入对东北土壤碳蓄积氮素利用效率的影响

由于人类活动影响,通过沉降和施肥方式进入生态系统的活性氮显著增加,其对土壤有机碳库产生重要影响。氮素利用效率(NUE)作为深入理解陆地生态系统碳氮耦合关系的重要参数,对NUE时空规律的研究不仅可以评估目前氮输入对陆地生态系统碳汇增加的贡献,同时也有助于预测未来氮输入情况下陆地生态系统的碳平衡。利用生态系统过程模型——CEVSA2模型的模拟结果,分析了东北地区氮输入情况下,土壤碳的氮素利用效率(SNUE)的时空变化规律及其影响因素,结果表明:(1)1961—2010年,氮输入的显著增加促进了土壤碳的蓄积,但SNUE显著下降;(2)森林的平均SNUE最高,农田最低;灌丛的下降速率最大,森林的SNUE变化趋势最不显著;(3)三江平原和长白山地区以及大小兴安岭的部分地区SNUE最大,其次是辽河平原、松嫩平原地区;内蒙古高原、呼伦贝尔高原地区以及大、小兴安岭的部分地区SNUE出现负值,说明在这些地区,外援氮输入抑制了土壤碳的蓄积;(4)氮输入的空间分异和不同生态系统响应氮输入的差异共同决定了SNUE及其变化的空间格局。该研究结果可为进一步分析不同区域氮促汇潜力和预测未来氮输入情景下的区域碳平衡提供参考。     

土壤氮磷添加下豆科草本植物生物固氮与磷获取策略的权衡机制

豆科草本植物固氮是陆地生态系统重要的自然氮输入方式, 影响着草地生产的经济性和可持续性。为探讨氮磷交互作用影响豆科草本植物生物固氮率的潜在生理生态机制, 该研究选取8种豆科草本植物分别种植在对照、氮肥添加、磷肥添加和氮磷耦合添加处理的土壤中, 进行野外盆栽实验。测定了初花期植物生物量和营养含量、根部碳水化合物含量、根际pH、根际柠檬酸含量、根际有效磷含量、植物根瘤生物量、磷含量及其生物固氮率。主要结果: 依赖于豆科物种, 氮添加显著促进了豆科草本植物根际磷的活化, 降低了根生物量分配以及根系非结构性碳水化合物含量。在两种磷添加处理下, 氮添加导致8种豆科草本植物根瘤生物量平均下降27%-36%, 生物固氮率平均下降20%-33%。磷添加降低了根际的磷活化, 但促进了豆科草本植物根系发育和非结构性碳水化合物的积累。在施氮和不施氮条件下, 磷添加分别使8种豆科草本植物的生物固氮率提高了45%-69%和0-47%。氮添加降低豆科草本植物生物固氮率, 其原因是氮添加提高了植物磷需求, 为活化更多磷, 豆科草本植物降低根系生物量和根系非结构性碳水化合物的含量, 导致根瘤发育受到限制。在氮添加的同时进行磷添加, 能够改善土壤氮磷平衡, 促进根系生长和非结构性碳水化合物积累, 缓解了增氮对生物固氮的抑制作用。     

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.     

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.     

Using indirect methods to constrain symbiotic nitrogen fixation rates: a case study from an Amazonian rain forest

Human activities have profoundly altered the global nitrogen (N) cycle. Increases in anthropogenic N have had multiple effects on the atmosphere, on terrestrial, freshwater and marine ecosystems, and even on human health. Unfortunately, methodological limitations challenge our ability to directly measure natural N inputs via biological N fixation (BNF)—the largest natural source of new N to ecosystems. This confounds efforts to quantify the extent of anthropogenic perturbation to the N cycle. To address this gap, we used a pair of indirect methods—analytical modeling and N balance—to generate independent estimates of BNF in a presumed hotspot of N fixation, a tropical rain forest site in central Rondônia in the Brazilian Amazon Basin. Our objectives were to attempt to constrain symbiotic N fixation rates in this site using indirect methods, and to assess strengths and weaknesses of this approach by looking for areas of convergence and disagreement between the estimates. This approach yielded two remarkably similar estimates of N fixation. However, when compared to a previously published bottom-up estimate, our analysis indicated much lower N inputs via symbiotic BNF in the Rondônia site than has been suggested for the tropics as a whole. This discrepancy may reflect errors associated with extrapolating bottom-up fluxes from plot-scale measures, those resulting from the indirect analyses, and/or the relatively low abundance of legumes at the Rondônia site. While indirect methods have some limitations, we suggest that until the technological challenges of directly measuring N fixation are overcome, integrated approaches that employ a combination of model-generated and empirically-derived data offer a promising way of constraining N inputs via BNF in natural ecosystems.     

Sources,fates, and impacts of nitrogen inputs to terrestrial ecosystems: review and synthesis

The relative importance of nitrogen inputs from atmospheric deposition and biological fixation is reviewed in a number of diverse, non-agricultural terrestrial ecosystems. Bulk precipitation inputs of N (l–l2 kg N ha^–1 yr^–1) are the same order of magnitude as, or frequently larger than, the usual range of inputs from nonsymbiotic fixation (< 1=" –=" 5=" kg=" n=">^–1 yr^–1), especially in areas influenced by industrial activity. Bulk precipitation measurements may underestimate total atmospheric deposition by 30–40% because they generally do not include all forms of wet and dry deposition. Symbiotic fixation generally ranges from 10–160 kg N ha^–1 yr^–1) in ecosystems where N-fixing species are present during early successional stages, and may exceed the range under unusual conditions.Rates of both symbiotic and nonsymbiotic fixation appear to be greater during early successional stages of forest development, where they have major impacts on nitrogen dynamics and ecosystem productivity. Fates and impacts of these nitrogen inputs are important considerations that are inadequately understood. These input processes are highly variable in space and time, and few sites have adequate comparative information on both nitrogen deposition and fixation.

A review on the effects of nitrogen and phosphorus addition on tree growth and productivity in forest ecosystems

Nitrogen (N) and phosphorus (P) inputs induced by anthropogenic activities and atmospheric N and P deposition have largely increased the availability of soil N and P in terrestrial ecosystems, which have considerably affected terrestrial carbon cycling processes. Tree growth and productivity in forest ecosystems play an important role in global carbon cycling, and determine the magnitude and direction of terrestrial carbon sequestration. Currently, a large number of field manipulation experiments have been conducted to investigate the effects of N and/or P addition on tree growth and forest productivity, but the results from these studies were inconsistent. Such inconsistent results might be affected by multiple factors, including biological, environmental and experimental variables. Here, we reviewed the present research status of the effects of N and P addition on tree growth and forest productivity in forest ecosystems based on three aspects, including the number of publications and experiments with field N and P addition, and the global distributions of these experiments. Then, we summarized the methods for assessing tree growth and forest productivity at ecosystem level in forest ecosystems, including relative growth rate and absolute increment. According to the related results, we reviewed the regulating factors that affect tree growth and productivity, and the potential mechanisms for such factors, including climate, tree size and stand age, plant functional traits (including type of tree-associated mycorrhizal fungi, N-fixation property of trees, and conservative and acquisitive functional traits), plant-microbe interaction, ambient nutrient (i.e., N and P) deposition rate, and experimental variables. Finally, we summarized the current studies, and pointed out five aspects that are urgently needed to provide further insights in future studies, including the physiological mechanism of how tree growth responds to N and P addition, the tradeoff and allocation among growth of various parts of tree under N and P addition, the role of plant functional traits in regulating and predicting the responses of tree growth to N and P addition, how the competition among trees regulates the responses of tree growth to N and P addition, and conducting long-term and coordinated distributed field experiments investigating the effects of N and P addition on tree growth and forest productivity at the global scale.     

Legume species identity and soil nitrogen supply determine symbiotic nitrogen-fixation responses to elevated atmospheric [CO2

In nitrogen (N)-limited systems, the response of symbiotic N fixation to elevated atmospheric [CO2] may be an important determinant of ecosystem responses to this global change. Experimental tests of the effects of elevated [CO2] have not been consistent. Although rarely tested, differences among legume species and N supply may be important. In a field free-air CO2 enrichment (FACE) experiment, we determined, for four legume species, whether the effects of elevated atmospheric [CO2] on symbiotic N fixation depended on soil N availability or species identity. Natural abundance and pool-dilution 15N methods were used to estimate N fixation. Although N addition did, in general, decrease N fixation, contrary to theoretical predictions, elevated [CO2] did not universally increase N fixation. Rather, the effect of elevated [CO2] on N fixation was positive, neutral or negative, depending on the species and N addition. Our results suggest that legume species identity and N supply are critical factors in determining symbiotic N-fixation responses to increased atmospheric [CO2].     

Relationships among phosphorus, molybdenum and free-living nitrogen fixation in tropical rain forests: results from observational and experimental analyses

Biological nitrogen (N) fixation is the primary source of “new” N to unmanaged ecosystems, and recent analyses suggest that terrestrial N inputs via free-living N fixation may be more important than previously assumed. This may be particularly true in some tropical rain forests, where free-living fixation could outpace symbiotic N fixation to represent the dominant source of new N inputs. However, our understanding of the controls over free-living N fixation in tropical rain forests remains poor, which directly constrains our ability to predict how N cycling will respond to changing environmental conditions. Although both phosphorus (P) and molybdenum (Mo) availability have been shown to limit free-living N fixation rates in the tropics, few studies have simultaneously explored P versus Mo limitation or the potential importance of P × Mo interactions. Here, an archived set of foliar, litter, and soil samples from a Costa Rican tropical rain forest provided an opportunity to simultaneously assess the relative strength of P versus Mo relationships with free-living N fixation rates. We also conducted a short-term, full-factorial (P × Mo) litter incubation experiment to directly assess nutrient limitation, allowing us to explore P and Mo controls over free-living N fixation rates using both observational and experimental approaches. We previously showed that N fixation rates were positively correlated with P concentrations in all substrates and, using the archived samples, we now show that Mo concentrations correlated with N fixation only in canopy leaves (where total Mo concentrations were extremely low). Likewise, fertilization with P alone (and not Mo) stimulated leaf litter N fixation rates. Thus, our results suggest that P availability dominantly controls free-living N fixation at this site, and when taken with data from other studies, our results suggest that attempts to identify “the nutrient” that limits N fixation in “the tropics” may be misguided. Rather, nutrient controls over free-living N fixation appear to be more nuanced—and the true nature of nutrient limitation to N fixation likely varies over a variety of scales across the vast tropical rain forest biome.     

Nitrogen deposition in tropical forests from savanna and deforestation fires

We used satellite‐derived estimates of global fire emissions and a chemical transport model to estimate atmospheric nitrogen (N) fluxes from savanna and deforestation fires in tropical ecosystems. N emissions and reactive N deposition led to a net transport of N equatorward, from savannas and areas undergoing deforestation to tropical forests. Deposition of fire‐emitted N in savannas was only 26% of emissions – indicating a net export from this biome. On average, net N loss from fires (the sum of emissions and deposition) was equivalent to approximately 22% of biological N fixation (BNF) in savannas (4.0 kg N ha^?1 yr^?1) and 38% of BNF in ecosystems at the deforestation frontier (9.3 kg N ha^?1 yr^?1). Net N gains from fires occurred in interior tropical forests at a rate equivalent to 3% of their BNF (0.8 kg N ha^?1 yr^?1). This percentage was highest for African tropical forests in the Congo Basin (15%; 3.4 kg N ha^?1 yr^?1) owing to equatorward transport from frequently burning savannas north and south of the basin. These results provide evidence for cross‐biome atmospheric fluxes of N that may help to sustain productivity in some tropical forest ecosystems on millennial timescales. Anthropogenic fires associated with slash and burn agriculture and deforestation in the southern part of the Amazon Basin and across Southeast Asia have substantially increased N deposition in these regions in recent decades and may contribute to increased rates of carbon accumulation in secondary forests and other N‐limited ecosystems.     

Biological Nitrogen Fixation in Two Tropical Forests: Ecosystem-Level Patterns and Effects of Nitrogen Fertilization

Humid tropical forests are often characterized by large nitrogen (N) pools, and are known to have large potential N losses. Although rarely measured, tropical forests likely maintain considerable biological N fixation (BNF) to balance N losses. We estimated inputs of N via BNF by free-living microbes for two tropical forests in Puerto Rico, and assessed the response to increased N availability using an on-going N fertilization experiment. Nitrogenase activity was measured across forest strata, including the soil, forest floor, mosses, canopy epiphylls, and lichens using acetylene (C₂H₂) reduction assays. BNF varied significantly among ecosystem compartments in both forests. Mosses had the highest rates of nitrogenase activity per gram of sample, with 11 ± 6 nmol C₂H₂ reduced/g dry weight/h (mean ± SE) in a lower elevation forest, and 6 ± 1 nmol C₂H₂/g/h in an upper elevation forest. We calculated potential N fluxes via BNF to each forest compartment using surveys of standing stocks. Soils and mosses provided the largest potential inputs of N via BNF to these ecosystems. Summing all components, total background BNF inputs were 120 ± 29 μg N/m²/h in the lower elevation forest, and 95 ± 15 μg N/m²/h in the upper elevation forest, with added N significantly suppressing BNF in soils and forest floor. Moisture content was significantly positively correlated with BNF rates for soils and the forest floor. We conclude that BNF is an active biological process across forest strata for these tropical forests, and is likely to be sensitive to increases in N deposition in tropical regions.