生态系统碳氮磷元素的生态化学计量学特征

生态系统元素平衡是当前全球变化生态学和生物地球化学循环的研究热点和焦点.在系统介绍生态化学计量学与碳氮磷元素循环研究进展的基础上,重点从土壤C:N:P化学计量比的分布特征、指示作用、对碳固定的影响,以及人类活动对C:N:P比的影响等方面探讨了C:N:P比在养分限制、生物地球化学循环、森林演替与退化等领域中的应用等问题,并展望了生态系统碳氮磷平衡的元素化学计量学未来研究的发展方向.通过对生态化学计量学理论和方法的研究,可以深入认识植物-凋落物-土壤相互作用的养分调控因素,对于揭示碳氮磷元素之间的相互作用及平衡制约关系,为减缓温室效应提供新思路和理论依据,具有重要的现实意义.     

樟子松人工林原产地与不同自然降水梯度引种地土壤和植物叶片生态化学计量特征

降水格局是影响陆地生态系统结构和过程的重要环境要素,尤其对于干旱/半干旱地区,降水变化是植物生长驱动的关键生态因子。目前,针对降水变化对陆地生态系统C、N、P等元素生物地球化学循环过程影响开展了大量研究。然而,关于沙地樟子松重要引种地科尔沁沙地自然降水梯度下沙地樟子松人工林土壤、植物生态化学计量特征的研究未见报道。因此,本研究以樟子松原产地红花尔基和引种地科尔沁沙地自然降水梯度下4个典型沙地樟子松人工林为对象,研究樟子松引种地降水变化对土壤(0—10,10—20 cm和20—40 cm)和植物(1年和2年生叶)生态化学计量特征的影响。研究结果发现:(1)与红花尔基原产地樟子松人工林相比,科尔沁沙地引种的樟子松人工林土壤C、N、P元素含量显著降低;(2)科尔沁沙地自西向东,随降水量增加,沙地樟子松人工林土壤C、N、P含量以及C∶P和N∶P表现为逐渐增加趋势,而土壤C∶N呈减少趋势;(3)随着降水量增加,樟子松叶C含量呈下降趋势,叶N含量和N∶P比值呈增加趋势,植物叶P含量无一致性规律;(4)樟子松叶片P含量与土壤C、N、P含量呈极显著正相关关系,而叶片C和N含量与土壤C、N、P含量无显著相关性。研究表明,沙地樟子松引种地科尔沁沙地土壤C、N、P养分比较缺乏,且随着降水增加土壤N养分限制降低,而土壤P养分限制增加。本研究从生态化学计量特征角度,为今后开展科尔沁沙地不同降水梯度条件下引种樟子松人工林提供理论依据。     

新疆67种荒漠植物叶碳氮磷计量特征及其与气候的关系

荒漠生态系统具有独特的耐旱植物种类和植物化学计量特征.本研究通过系统采集全疆63个荒漠地点的67种植物,探索荒漠植物叶碳、氮、磷(C、N、P)的计量特征及其与气候因子的关系.结果表明:荒漠植物叶C、N和P的平均含量分别为394、18.4和1.14 mg·g-1,C∶N、C∶P和N∶P平均值分别为28、419和18.灌木的N含量高于乔木和草本,灌木P含量比草本(乔木)低(高);C3植物叶的C、N、C∶P、N∶P高于C4植物.随年均降水量增加,叶C有先降低后升高的趋势,叶N、P先升高、后降低,叶C∶N、C∶P与叶N、P变化趋势相反,N∶P变化不显著;随年均温升高,叶C先降低后升高,叶N、P降低,C∶N变化不显著,C∶P、N∶P升高.年均降水量对叶片化学计量特征的影响大于年均温度和植物类型.本研究结果将有助于预测全球气候变化背景下的C、N、P元素循环规律的变化,并为干旱区生物地球化学建模提供参考和基础数据.     

中国典型生态脆弱区生态化学计量学研究进展

在人类活动和自然环境变化的相互作用下,生态脆弱区生态系统随之变迁,荒漠化、盐碱化、水土流失、植被生产力下降等是生态脆弱区面临的重要问题。生态化学计量学作为当前多学科交叉研究的热点领域,强调从生态系统能量与元素平衡角度,揭示元素生物地球化学循环和生态系统对环境变化的调控机制。为了促进对生态脆弱区碳（C）、氮（N）、磷（P）生态化学计量的深入理解,本文重点总结了近年来有关我国典型生态脆弱区植物、凋落物、土壤和土壤微生物量C、N、P生态化学计量及其对环境变化响应的研究成果,并展望未来研究方向,以期促进生态化学计量学的发展和生态脆弱区生态保护与恢复研究。研究表明,植物-凋落物-土壤-土壤微生物系统C、N、P化学计量具有较强相关性,并受土壤因子、气候因子、生物因子和人类活动的显著影响。在生态脆弱区,我国北方荒漠及荒漠化地区由于较高的N∶P比值易受P限制,而青藏高原脆弱区、西南岩溶石漠化地区和黄土高原脆弱区等生态脆弱区更易受N限制;随着植被恢复,养分限制逐渐由N限制向P限制转变。生态脆弱区相对较低的养分含量和C∶N∶P比值或许可在一定程度上解释植被生产力较低的原因,而具有较高N、P化学计量内稳性的植物在贫瘠环境中具有较强竞争力和更高稳定性。今后可加强多尺度、不同生态系统植物-凋落物-土壤-土壤微生物系统生态化学计量和长期、多因子交互控制实验的研究。     

土壤微生物生物量碳氮磷与土壤酶化学计量对气候变化的响应机制

微生物和土壤酶是陆地生态系统中生物地球化学循环的重要驱动力,深入理解微生物在生态系统中的调节作用以及气候变化过程中微生物量和土壤酶的响应机制是生态学领域关注的重要科学问题.本研究从气候因素角度出发,基于生态化学计量学理论,综述了微生物和土壤酶在陆地生态系统碳氮磷循环中的作用,以及土壤微生物生物量碳氮磷和土壤酶化学计量对气候变化的响应机制,即: 改变微生物代谢速率和酶活性;调整微生物群落结构;调整微生物生物量碳氮磷与土壤酶化学计量特征;改变碳氮磷养分元素利用效率.最后分析当前研究的不足,并提出了该领域亟待解决的科学问题: 综合阐明土壤微生物和土壤酶对气候变化的响应机制;探究土壤微生物和胞外酶养分耦合机理;深入探究土壤微生物量和土壤酶化学计量特征对气候变化的适应对策.     

荒漠生态系统磷循环及其驱动机制研究进展

磷(P)循环在维持荒漠生态系统的生物多样性水平、结构和功能的稳定性、元素的动态平衡,以及荒漠自然资源的可持续利用方面有重要作用。通过查阅国内外有关P循环的文献资料,发现当前国内尚缺乏针对荒漠生态系统P循环的系统研究,特别是P循环的生物和非生物驱动机制。综述了荒漠生态系统P的输入-输出过程,植物对P的吸收转运机制以及对P循环的作用,生物土壤结皮(BSC)有机分泌物对P循环的贡献,以及荒漠生态系统P循环过程对气候变化的响应机制等。文末展望了荒漠生态系统P循环的一些重要研究方向和亟需解决的科学问题,包括(1)P在荒漠生态系统中的存在形态、分配及动态平衡;(2)土壤微生物对荒漠植物获取土壤有效P的驱动作用;(3)入侵植物对P循环的影响与潜在生态风险评估;(4)利用分子生物学和基因组学手段揭示真菌-荒漠植物根系系统P循环的基因调控机制;(5)微生物分泌物、土壤磷酸酶类(包括磷酸单酯酶、磷酸二酯酶和三磷酸单酯水解酶)和作用于含磷酸酐和N—P键的酶对土壤P循环的调控;(6)气候变化(干旱、高温和降水节律变化等)如何影响P的生物和非生物转化过程;(7)基于同位素示踪和生态化学计量学理论解释荒漠生态系统...     

元谋干热河谷燥红土和变性土上植物叶片的元素含量及其重吸收效率

采用盆栽试验,研究元谋干热河谷燥红土和变性土上生长的植物叶片以及凋落叶营养元素含量,并分析养分重吸收效率对土壤类型与物种互作的响应.结果表明: 土壤类型对叶片N、P、Ca、Mg、Cu、Zn、Fe、N∶P以及凋落叶N、P、Mn、N∶P均有显著影响;燥红土植物叶片与凋落叶N、Mn含量和N∶P显著高于变性土,而燥红土植物叶片P、Ca、Mg、Fe、Cu、Zn和凋落叶P含量显著低于变性土.燥红土植物叶片N含量较变性土高34.8%,而P含量低40.0%;在叶片凋落时,N、P、K表现为重吸收,而其他元素呈富集状态.燥红土凋落叶Ca、Mg、Mn富集系数显著高于变性土.物种仅对叶片N含量有显著影响,物种与土壤交互作用对植物叶片和凋落叶元素含量影响不显著,表明各土壤类型对不同物种元素含量的影响方式较为一致.土壤类型对植物元素含量的影响可进一步作用于干热河谷植物凋落物分解、植物-土壤的养分反馈以及生物地球化学循环.     

氮输入对植物光合固碳的影响研究进展

植物光合固碳(C)是生物固C的重要途径和生态系统C循环中的重要环节。在全球环境变化背景下,研究氮(N)输入对植物光合固C的影响,对于更好的认识C、N循环过程及生态系统对全球变化的响应过程等具有重要意义。N输入是否能够增加植物固C取决于生态系统类型以及生态系统的N饱和度;草原和湿地生态系统N输入的临界负荷值较高,干旱、半干旱荒漠地区较低;N输入可能改变植物光合固C在各器官的分配,主要由植物生理、自身生长节律和环境养分等决定。由于物种和生态系统类型的差异,N输入对植物固C的影响仍具有很大的不确定性,目前缺乏准确、定量表达N输入对生态系统光合和C同化物分配影响的数学表达方法和过程算法。未来应着重加强N输入下C同化物分配的生物地球化学模型和N、P富集下植物光合固C耦合模型研究,并应用同位素标记和分子生物学技术,从生态系统角度综合探讨N输入下植物光合固C的分配和转化特征。     

季节性雪被变化对森林凋落物分解及土壤氮动态的影响

全球气候变化引发的雪被格局变化将深刻影响植被的凋落物分解、陆地生态系统的土壤养分循环等过程.森林是陆地生态系统的主体,在全球生物地球化学循环中起着不可替代的作用.本研究综述了季节性雪被变化对森林凋落物分解及土壤氮动态的影响.全球气候变化情景下季节性雪被表现出因地域而异的增加或减少的变化格局,一方面通过改变环境温湿度、凋落物质量、分解者动态等直接影响分解过程,另一方面通过改变森林群落结构、植被物候、土壤养分等间接地作用于凋落物分解.同时,季节性雪被通过影响氮富集作用、雪被下土壤温湿度、冻融循环、森林群落、雪下动物和微生物等相关因子而改变森林土壤氮循环.本领域未来应开展的研究是: 1) 全面考虑全球气候变化情景下季节性雪被格局的变异性,开展不同季节性雪被格局变化的模拟研究;2) 开展季节性雪被融雪水淋溶作用对森林凋落物分解和土壤氮动态的影响研究;3) 阐明不同生态系统和气候带中季节性雪被格局变化对森林凋落物分解过程和土壤氮动态的驱动机制研究;4) 量化季节性雪被变化对森林凋落物分解和土壤氮动态在雪被覆盖期的瞬时影响和无雪期的延续影响,为阐明和模型预测陆地生态系统生物地球化学循环对全球气候变化的响应提供理论基础和数据支持.     

土壤微生物对气候变暖和大气N沉降的响应

气候变暖和大气N沉降是近一、二十年来人们非常关注的全球变化现象,它们所带来的一系列生态问题已成为全球变化研究的重要议题。它们不仅影响地上植被生长和群落组成,还直接或间接地影响土壤微生物过程,而土壤微生物对此做出的响应正是生态系统反馈过程中非常重要的环节。该文分别从气候变化对土壤微生物的影响(土壤微生物量、微生物活动和微生物群落结构)和土壤微生物对气候变化的响应(凋落物分解、养分利用与循环以及养分的固持与流失)两个角度,综述近期土壤微生物对气候变暖和大气N沉降响应与适应的研究进展。气候变暖和大气N沉降对土壤微生物的影响更多地反映在微生物群落的结构和功能上,而土壤微生物量、微生物活动和群落结构的变化又会通过改变凋落物分解、养分利用和C、N循环等重要的土壤生态系统功能和过程做出响应,形成正向或负向反馈,加强或削弱气候变化给整个陆地生态系统带来的影响。然而,到目前为止土壤微生物的响应对陆地生态系统产生的最终结果仍是未决的关键性问题。     

陆地生态系统植被氮磷化学计量研究进展

因化学功能的耦合和元素的不可替代性, 植物对N、P的需求和利用存在严格的比例。植物N、P化学计量在不同功能群、生长地区、生长季、器官之间以及环境梯度下存在明显的变化规律。多数研究从N、P浓度、N:P及N、P间异速指数等角度分析了植物化学计量变化规律, 并探讨其在全球范围内的具体数值。为增进对植物响应全球变化的理解, 该文综述了N、P化学计量的影响因素及其机理的最新研究进展, 并指出未来拟重点研究的方向。     

Are patterns in nutrient limitation belowground consistent with those aboveground: results from a 4 million year chronosequence

Accurately predicting the effects of global change on net carbon (C) exchange between terrestrial ecosystems and the atmosphere requires a more complete understanding of how nutrient availability regulates both plant growth and heterotrophic soil respiration. Models of soil development suggest that the nature of nutrient limitation changes over the course of ecosystem development, transitioning from nitrogen (N) limitation in ‘young’ sites to phosphorus (P) limitation in ‘old’ sites. However, previous research has focused primarily on plant responses to added nutrients, and the applicability of nutrient limitation-soil development models to belowground processes has not been thoroughly investigated. Here, we assessed the effects of nutrients on soil C cycling in three different forests that occupy a 4 million year substrate age chronosequence where tree growth is N limited at the youngest site, co-limited by N and P at the intermediate-aged site, and P limited at the oldest site. Our goal was to use short-term laboratory soil C manipulations (using ¹⁴C-labeled substrates) and longer-term intact soil core incubations to compare belowground responses to fertilization with aboveground patterns. When nutrients were applied with labile C (sucrose), patterns of microbial nutrient limitation were similar to plant patterns: microbial activity was limited more by N than by P in the young site, and P was more limiting than N in the old site. However, in the absence of C additions, increased respiration of native soil organic matter only occurred with simultaneous additions of N and P. Taken together, these data suggest that altered nutrient inputs into ecosystems could have dissimilar effects on C cycling above- and belowground, that nutrients may differentially affect of the fate of different soil C pools, and that future changes to the net C balance of terrestrial ecosystems will be partially regulated by soil nutrient status.     

Temporal changes in soil C‐N‐P stoichiometry over the past 60 years across subtropical China

Controlled experiments have shown that global changes decouple the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), resulting in shifting stoichiometry that lies at the core of ecosystem functioning. However, the response of soil stoichiometry to global changes in natural ecosystems with different soil depths, vegetation types, and climate gradients remains poorly understood. Based on 2,736 observations along soil profiles of 0–150 cm depth from 1955 to 2016, we evaluated the temporal changes in soil C‐N‐P stoichiometry across subtropical China, where soils are P‐impoverished, with diverse vegetation, soil, and parent material types and a wide range of climate gradients. We found a significant overall increase in soil total C concentration and a decrease in soil total P concentration, resulting in increasing soil C:P and N:P ratios during the past 60 years across all soil depths. Although average soil N concentration did not change, soil C:N increased in topsoil while decreasing in deeper soil. The temporal trends in soil C‐N‐P stoichiometry differed among vegetation, soil, parent material types, and spatial climate variations, with significantly increased C:P and N:P ratios for evergreen broadleaf forest and highly weathered Ultisols, and more pronounced temporal changes in soil C:N, N:P, and C:P ratios at low elevations. Our sensitivity analysis suggests that the temporal changes in soil stoichiometry resulted from elevated N deposition, rising atmospheric CO₂ concentration and regional warming. Our findings revealed that the responses of soil C‐N‐P and stoichiometry to long‐term global changes have occurred across the whole soil depth in subtropical China and the magnitudes of the changes in soil stoichiometry are dependent on vegetation types, soil types, and spatial climate variations.     

Effects of three global change drivers on terrestrial C:N:P stoichiometry: a global synthesis

Over the last few decades, there has been an increasing number of controlled‐manipulative experiments to investigate how plants and soils might respond to global change. These experiments typically examined the effects of each of three global change drivers [i.e., nitrogen (N) deposition, warming, and elevated CO₂] on primary productivity and on the biogeochemistry of carbon (C), N, and phosphorus (P) across different terrestrial ecosystems. Here, we capitalize on this large amount of information by performing a comprehensive meta‐analysis (>2000 case studies worldwide) to address how C:N:P stoichiometry of plants, soils, and soil microbial biomass might respond to individual vs. combined effects of the three global change drivers. Our results show that (i) individual effects of N addition and elevated CO₂ on C:N:P stoichiometry are stronger than warming, (ii) combined effects of pairs of global change drivers (e.g., N addition + elevated CO₂, warming + elevated CO₂) on C:N:P stoichiometry were generally weaker than the individual effects of each of these drivers, (iii) additive interactions (i.e., when combined effects are equal to or not significantly different from the sum of individual effects) were more common than synergistic or antagonistic interactions, (iv) C:N:P stoichiometry of soil and soil microbial biomass shows high homeostasis under global change manipulations, and (v) C:N:P responses to global change are strongly affected by ecosystem type, local climate, and experimental conditions. Our study is one of the first to compare individual vs. combined effects of the three global change drivers on terrestrial C:N:P ratios using a large set of data. To further improve our understanding of how ecosystems might respond to future global change, long‐term ecosystem‐scale studies testing multifactor effects on plants and soils are urgently required across different world regions.     

土壤有效氮及其相关因素对植物细根的影响

细根(直径≤2mm)作为植物吸收水分和养分的主要器官之一,在陆地生态系统养分循环和能量流动中起重要作用。开展土壤有效氮变化对植物细根影响研究对于了解全球气候变化条件下的陆地生态系统养分循环具有重要意义。本文就相关研究进行了综述:1)土壤有效氮变化对植物细根生长、发育、寿命及呼吸的直接影响;2)土壤质地、温度、大气CO2浓度和氮沉积等相关因素对植物细根的影响。由于研究方法及物种间差异等的影响,研究结果不尽相同。今后,应在不同空间尺度上深入研究土壤有效氮对植物细根的影响,而植物细根-土壤-微生物三者间相互关系变化对土壤氮变化的潜在响应将可能成为今后研究的热点问题之一。     

草地植物根系碳储量和碳流转对CO2浓度升高的响应

植物根系是陆地生态系统重要的碳汇和矿质养分库,也是土壤中碳及养分的主要来源,只有深入认识CO2浓度升高下根系的碳汇功能和根系周转对土壤碳库的影响,才能准确预测生态系统对全球变化的响应与反馈调节作用.介绍了CO2浓度升高对草地植物根系生物量、根系凋落物的数量和品质以及根系周转速率的影响,指出研究植物体内碳向根分配格局的变化趋势必须考虑CO2浓度升高的直接和间接两方面作用;在预测根系碳库储量的动态变化时,需要区分不同功能根系组分的生物量;为更准确估算根系周转速率,有必要确立草地植物根系直径与其寿命之间的关系;CO2浓度升高普遍提高根系凋落物的C/N,但以此判定其在土壤中的分解速率快慢并不可靠,需要进一步从机理上探究根系凋落物分解的控制因素.     

A Synthesis of Climate and Vegetation Cover Effects on Biogeochemical Cycling in Shrub-Dominated Drylands

Semi-arid and arid ecosystems dominated by shrubs (“dry shrublands”) are an important component of the global C cycle, but impacts of climate change and elevated atmospheric CO₂ on biogeochemical cycling in these ecosystems have not been synthetically assessed. This study synthesizes data from manipulative studies and from studies contrasting ecosystem processes in different vegetation microsites (that is, shrub or herbaceous canopy versus intercanopy microsites), to assess how changes in climate and atmospheric CO₂ affect biogeochemical cycles by altering plant and microbial physiology and ecosystem structure. Further, we explore how ecosystem structure impacts on biogeochemical cycles differ across a climate gradient. We found that: (1) our ability to project ecological responses to changes in climate and atmospheric CO₂ is limited by a dearth of manipulative studies, and by a lack of measurements in those studies that can explain biogeochemical changes, (2) changes in ecosystem structure will impact biogeochemical cycling, with decreasing pools and fluxes of C and N if vegetation canopy microsites were to decline, and (3) differences in biogeochemical cycling between microsites are predictable with a simple aridity index (MAP/MAT), where the relative difference in pools and fluxes of C and N between vegetation canopy and intercanopy microsites is positively correlated with aridity. We conclude that if climate change alters ecosystem structure, it will strongly impact biogeochemical cycles, with increasing aridity leading to greater heterogeneity in biogeochemical cycling among microsites. Additional long-term manipulative experiments situated across dry shrublands are required to better predict climate change impacts on biogeochemical cycling in deserts.