氮添加对中国陆地植被地上-地下生物量分配的影响

大气氮沉降水平持续升高导致的外源氮输入增加,强烈影响了陆地生态系统的碳循环。目前,已有大量报道证实了氮沉降升高对全球陆地植被固碳的积极影响。虽然之前大部分研究将这一结果归因于光合作用增强导致的地上生物量增加,但最近的研究发现长期氮添加对植物地下根系的影响也同样重要。归纳整理了181篇公开发表的我国野外模拟氮沉降试验结果,采用整合分析(Meta-analysis)方法,定量评估了氮添加对我国陆地植被地上-地下生物量分配的影响特征和不同生态系统类型及施氮方式之间的影响差异。通过分析地上-地下生物量分配对氮添加的响应差异来探究植被碳增益对长期大气氮沉降增加的潜在响应机制。结果表明,氮添加显著增强了我国陆地植被的光合作用及碳固存,且植物碳增益在不同生态系统类型及施氮制度间有所差异。植物叶片的氮含量显著增加,使得叶片碳氮比及凋落物碳氮比显著降低,但并未显著影响细根的碳氮比。氮添加总体上显著提高了植物的净光合速率,但降低了光合利用效率。地上生物量,凋落物产量和根生物量平均分别显著增加了38%,17%和18%,总体上植物地上部分对氮添加的响应程度比地下部分更高。然而,不同生态系统类型的地上-地下生物...     

土壤有机碳对外源氮添加的响应及其机制

土壤有机碳库是陆地生态系统碳库的重要组成, 在全球碳循环中发挥着重要的作用。受元素化学计量平衡调控作用, 氮输入的增加将会对土壤有机碳库产生重要影响。然而, 目前关于陆地生态系统碳库对氮添加的响应主要集中在植被碳库, 对土壤碳库研究较少, 且研究结论争议较大, 尤其对其响应机制缺少系统梳理。该文作者通过对已有文献进行梳理, 认为生态系统类型、土壤碳变化的检测方法、土壤深度, 以及土壤稳定性碳和易变碳含量的差异可能是造成当前研究土壤碳汇增量(每克氮输入所增加的碳)差异的重要原因。氮添加条件下土壤有机碳的积累机制可能包括3个方面: 1)氮添加增加了凋落物输入, 促进了碳积累; 2)氮添加减少土壤碳输出, 尤其是抑制了稳定性碳的分解; 3)促进土壤腐殖质及稳定性碳的形成。此外, 该文结合当前研究中存在的不足, 提出今后需加强对深层土壤碳、土壤可溶性有机碳的淋溶及吸附, 以及不同土壤碳组分对氮添加的响应研究, 并通过改进检测方法减少氮添加条件下碳储量的测量误差。     

土地利用方式对西双版纳热带森林土壤微生物生物量碳的影响

2006年1月至2007年9月,通过去除凋落物和切根控制试验,研究了热带森林不同土地利用方式(次生林/橡胶林)对西双版纳热带森林土壤微生物生物量碳的影响.结果表明:次生林转变为橡胶林后,土壤养分及植物碳输入均明显减少,土壤微生物生物量碳显著降低;两种林型雨季土壤微生物生物量碳均高于干季,其中次生林土壤微生物生物量碳与土壤温度呈显著正相关,而橡胶林则与土壤湿度呈正相关关系;次生林土壤微生物生物量碳受植物根系养分输入的调控,地上凋落物数量的影响较小,土壤微生物生物量碳与细根生物量及其C、N输入量呈显著正相关;橡胶林细根生物量及其C、N输入量,以及地上凋落物均未显著影响土壤微生物生物量碳.橡胶种植导致土壤养分含量和土壤pH值降低,加上严重的人为干扰,土壤微生物生物量碳减少,且其调控因素发生显著变化,最终可能影响土壤的其他生态过程.     

氮添加增加中国陆生植物生物量并降低其氮利用效率

研究植物整体和地上、地下部分生物量和氮利用效率(NUE)对氮添加的响应,有助于了解全球氮沉降背景下中国陆地生态系统植物碳、氮循环的变化。然而氮添加对植物生物量和NUE的影响是否一致,影响这种响应的主要因素仍不清楚。该研究收集整理了94篇已发表论文中有关中国氮添加实验的相关数据,使用meta分析定量评估了氮添加对植物生物量分配和NUE的影响及其主要影响因素。结果表明:(1)氮添加在显著增加植物整体、地上和地下生物量的同时,也显著降低了不同部分的NUE,但植物地上生物量的增加幅度(34.0%)高于地下生物量(5.3%),而地下部分NUE(29.9%)的下降幅度高于地上部分NUE (15.4%);(2)植物整体和不同部分生物量及其NUE对氮添加的响应在不同生态系统类型、氮肥形式、施氮水平、持续时间和水分条件下均存在显著差异,且草地和荒漠生态系统对氮添加的响应明显高于其他生态系统类型;(3)影响植物整体、地上和地下生物量效应值的最主要因素均为土壤总氮含量,植物整体和地上部分NUE的最主要影响因素均为施氮量,而影响地下部分NUE效应值的最主要因素为氮肥形式。总之,植物生物量和NUE在氮添加条件下...     

氮沉降增加情景下植物-土壤-微生物交互对自然生态系统土壤有机碳的调控研究进展

大气氮沉降增加倾向于促进受氮限制陆地生态系统地上生物量,但是对地下碳过程和土壤碳截存的影响结果迥异,导致陆地生态系统“氮促碳汇”的评估存在很大的不确定性。大气氮沉降输入直接影响微生物活性或间接影响底物质量,改变凋落物和土壤有机质(SOM)的分解速率和分解程度,进而影响土壤有机碳(SOC)的积累与损耗过程。过去相关研究主要集中在土壤碳转化过程和碳储量动态方面,缺乏植物-微生物-SOM交互作用的理解,对土壤碳截存调控的生物化学和微生物学机理尚不清楚。本文以地下碳循环过程为主线,分别综述了氮沉降增加对植物地下碳分配、SOC激发效应、微生物群落碳代谢过程的影响,深入分析SOM化学稳定性与微生物群落动态的关系。该领域研究的薄弱环节体现在:(1)增氮倾向于降低根系的生长和周转,对根际沉积碳分配(数量和格局)的影响及驱动因素不明确;(2)虽然认识到氮素有效性影响土壤激发效应的方向和强度,但是氧化态NO-3和还原态NH+4输入对有机质激发效应的差异性影响及潜在机理知之甚少;(3)微生物碳利用效率(CUE)是微生物群落碳代谢的关键表征,能够很好地解释土壤碳的积累与损耗过程;由于缺乏适宜的测定方法,难以准确量化土壤微生物的CUE及微生物生物量的周转时间;(4)增氮会抑制土壤真菌群落及其胞外酶活性,对细菌群落组成的影响尚未定论,有关SOM化学质量与土壤微生物群落活性、组成之间的耦合关系尚不清楚。未来研究应基于长期的氮添加控制实验平台,结合碳氧稳定性同位素示踪、有机质化学、分子生物学和宏基因组学等方法,深入分析植物同化碳的地下分配规律、微生物碳代谢和周转、有机质化学结构与功能微生物群落的耦合关系等关键环节。上述研究将有助于揭示植物-土壤-微生物交互作用对SOC动态的调控机制,完善陆地生态系统碳-氮耦合循环模型,有效降低区域陆地碳汇评估的不确定性,并可为陆地生态系统应对全球变化提供科学依据。     

陆地生态系统地下碳输入与输出过程研究进展

生态系统地下碳输入与输出过程是陆地生态系统碳分配和转化的核心,并直接影响着全球碳循环。陆地生态系统凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸是地下碳输入与输出过程中的重要组成部分。由于这些组分非常复杂且其研究技术和方法受到限制,目前人们对陆地生态系统地下碳输入与输出过程尚缺乏全面的认识,故在陆地生态系统碳循环研究中存在诸多的不确定性。该文概述了凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸的研究方法,以及它们对气候变化的响应,探讨了陆地生态系统地下碳输入与输出过程中的研究难点,并对未来需要深入探究的一些领域进行了展望。     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

外来植物入侵对陆地生态系统地下碳循环及碳库的影响

生物入侵是当今全球性重大环境问题之一, 是全球变化的主要研究内容.评价外来植物入侵对于生态系统影响的研究多集中在地上部分,对于生态系统地下部分影响的研究相对较少.陆地生态系统地下部分对于生态系统过程的重要性之一体现在它处于生态系统碳分配过程的核心环节.入侵种通过影响群落凋落物的输入数量、质量以及输入时间,影响到对于土壤的碳输入,而入侵种与土著种根系的差异以及入侵种对微生物群落的影响是造成土壤呼吸强度发生变化的主要因素,前者土壤呼吸强度一般比后者高.多数研究表明外来植物入侵对生态系统地下碳循环和碳库产生影响,但由于入侵植物种类较多以及研究地点环境条件的不同,关于外来植物入侵对于土壤碳库和土壤有机碳矿化影响的研究结论并不统一.最后,提出了今后该研究领域应加强的一些建议和方向.     

川西高原季节性雪被覆盖下凋落物输入对土壤微生物数量及生物量的影响

为了了解季节性雪被覆盖下不同碳供应水平对高山土壤生态系统过程的影响,2010 年1 月-5 月在青藏高原东缘设计人工雪厚度梯度控制(0 cm, 30 cm, 100 cm)和凋落物添加(0 g, 5 g, 20 g 鲜卑花叶片)的原位试验,测定了土壤中的微生物数量和微生物生物量。研究发现,雪被覆盖能有效地绝缘大气和土壤,减少冻融交替的幅度和频次,显著增加了细菌和真菌数量,而对微生物生物量碳氮无明显影响。凋落物的输入降低了微生物生物量氮的含量,增加了细菌和真菌的数量。说明雪被覆盖和有机碳的输入可以通过影响冬季土壤微生物群落结构,从而对高山地区冬季生态系统过程产生实质性的影响。     

植物群落向土壤有机碳输入及其对气候变暖的响应研究进展

植物群落作为陆地生态系统土壤有机碳的主要来源,可通过地表凋落物分解、细根周转和根系分泌物等方式将光合作用同化的碳输入到土壤中。全球气候变暖正深刻地影响植物群落的分布、结构与功能,改变森林地上和地下凋落物产量与分解速率和根系分泌过程,从而改变植物群落向土壤输入有机碳数量。本文综述了植物群落向土壤有机碳输入过程及其对气候变暖的响应研究进展。研究表明,气候变暖可通过影响植物群落生产直接影响凋落物产量和根系分泌过程,还可通过改变凋落物分解环境条件、凋落物基质质量和分解者群落结构与活性等非生物与生物因子而间接作用于凋落物向土壤有机碳输入过程。气候变暖还可通过影响植物根系性状、根系分泌物化学组成等间接影响植物根系向土壤输入的碳量,但其具体机制还需深入探讨。未来的研究应该关注气候变暖导致植物群落结构改变进而影响土壤有机碳输入的具体机制以及粗木质残体对土壤有机碳输入的贡献,同时还应注重植物凋落物与根系分泌过程的整合研究,以期更全面地认识气候变暖背景下植物群落对土壤碳库及碳循环过程的贡献。     

亚热带典型森林地上和地下凋落物输入对土壤新老有机碳动态平衡的影响

凋落物是森林土壤有机碳(SOC)形成、稳定和周转的重要影响因子。目前针对亚热带不同类型森林地上和地下凋落物对新SOC累积和老SOC输出动态平衡的影响仍不清楚。本研究以中亚热带常绿阔叶天然林、马尾松人工林和杉木人工林为对象,基于C3/C4植物-土壤置换试验,利用稳定同位素¹³C示踪方法开展3年野外定位试验,分析了森林地上、地下凋落物输入对SOC周转的影响。结果表明: 森林类型、凋落物处理和时间均能显著影响SOC含量、土壤δ¹³C值、新SOC和老SOC含量,且存在显著的森林类型×凋落物处理交互效应。地上和地下凋落物输入均能显著提高SOC含量和净增量,与杉木人工林相比,天然林SOC对凋落物输入的响应更敏感。凋落物输入显著降低了土壤δ¹³C值,且天然林、马尾松人工林土壤δ¹³C显著低于杉木人工林。在马尾松人工林,地下凋落物处理的新SOC含量显著高于地上凋落物;在天然林和马尾松人工林,地下凋落物输入处理的老SOC含量显著低于地上凋落物处理。此外,地上凋落物归还量和地下根生物量与SOC含量和净增量呈显著正相关,而地下根凋落物量和C/N与新SOC含量呈显著正相关。森林地下凋落物比地上凋落物输入对SOC周转的影响更重要,且不同森林凋落物输入对SOC的影响存在差异性。本研究可为揭示亚热带典型森林土壤有机碳库的形成和可持续管理提供依据。     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Can litter addition mediate plant productivity responses to increased precipitation and nitrogen deposition in a typical steppe?

Plant litter is a key component of grassland and plays a major role in terrestrial ecosystem processes. Global climate change has been shown to considerably alter litter inputs to soils, which may feed back to the grassland ecosystem responses to climate change. In order to explore whether litter addition could mediate above and belowground productivity responses to short-term increases in growing-season precipitation and nitrogen deposition, we conducted a two-year study on water, nitrogen and litter addition in Inner Mongolia grassland. After two years of treatments, our results showed that water, nitrogen, and litter addition increased aboveground biomass (AB) and belowground net primary productivity (BNPP). Besides, litter addition increased BNPP responses to water addition. These litter addition effects could be attributed to the influence of litter on soil moisture and soil nitrogen availability, ultimately increasing belowground water use efficiency (WUE_BNPP) and plant nitrogen uptake (NUP_BNPP). However, litter addition suppressed the aboveground biomass (AB) responses to nitrogen addition under ambient precipitation conditions by affecting soil moisture. In conclusion, our results suggest that ecosystem responses to short-term increases in growing-season precipitation and nitrogen deposition could be mediated by the increased litter input caused by climate change.     

酸雨对中国陆地生态系统土壤呼吸影响的整合分析

通过整合分析(Meta-analysis)国内外公开发表的81篇模拟酸雨实验论文的2683条有效观测数据,量化了酸雨对中国3个主要陆地生态系统(森林、草地和农田)土壤呼吸(Rs)及其组分(自养呼吸—Ra、异养呼吸—Rh)的影响。结果表明,酸雨显著降低了Rs(-9.6%)、Rh(-7.7%)和Ra(-11.7%);酸雨pH越低,Rs及其组分的降幅越大;野外实验对Rh和Ra的负效应大于温室实验。酸雨对Rs的负效应在农田最大(-14.7%),草地次之(-10.8%),森林最小(-8.0%);森林Rh、Ra对酸雨的响应一致,不同林型间差异不显著;草地Rh和Ra在酸雨处理下分别显著降低和增加。Rs、Rh与土壤pH显著正相关,与土壤有机碳(SOC)显著负相关;Rh和Ra分别与地上和地下生物量(AGB和BGB)显著正相关。酸雨对Rs和Ra的负效应随纬度的增加而减弱,随MAT的升高而增强,对Rs的正效应随MAP的降低而增强。本研究表明,酸雨不仅降低了土壤pH,抑制了植物生长,减少了植物向土壤的碳输入,还降低了微生物活性,减少了Rh,导致SOC分解降低,因而未显著改变土壤碳库。     

Hierarchical plant responses and diversity loss after nitrogen addition: testing three functionally-based hypotheses in the Inner Mongolia grassland

Background

Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done.

Methodology/Principal Findings

In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem.

Conclusions/Significance

Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

Nitrogen addition promotes terrestrial plants to allocate more biomass to aboveground organs: A global meta-analysis

A significant increase in reactive nitrogen (N) added to terrestrial ecosystems through agricultural fertilization or atmospheric deposition is considered to be one of the most widespread drivers of global change. Modifying biomass allocation is one primary strategy for maximizing plant growth rate, survival, and adaptability to various biotic and abiotic stresses. However, there is much uncertainty as to whether and how plant biomass allocation strategies change in response to increased N inputs in terrestrial ecosystems. Here, we synthesized 3516 paired observations of plant biomass and their components related to N additions across terrestrial ecosystems worldwide. Our meta-analysis reveals that N addition (ranging from 1.08 to 113.81 g m⁻² year⁻¹) increased terrestrial plant biomass by 55.6% on average. N addition has increased plant stem mass fraction, shoot mass fraction, and leaf mass fraction by 13.8%, 12.9%, and 13.4%, respectively, but with an associated decrease in plant reproductive mass (including flower and fruit biomass) fraction by 3.4%. We further documented a reduction in plant root-shoot ratio and root mass fraction by 27% (21.8%–32.1%) and 14.7% (11.6%–17.8%), respectively, in response to N addition. Meta-regression results showed that N addition effects on plant biomass were positively correlated with mean annual temperature, soil available phosphorus, soil total potassium, specific leaf area, and leaf area per plant. Nevertheless, they were negatively correlated with soil total N, leaf carbon/N ratio, leaf carbon and N content per leaf area, as well as the amount and duration of N addition. In summary, our meta-analysis suggests that N addition may alter terrestrial plant biomass allocation strategies, leading to more biomass being allocated to aboveground organs than belowground organs and growth versus reproductive trade-offs. At the global scale, leaf functional traits may dictate how plant species change their biomass allocation pattern in response to N addition.     

黄土区不同退耕方式下土壤碳氮的差异及其影响因素

研究植被恢复对土壤碳氮动态的影响,对了解陆地生态系统碳氮循环,应对全球温室效应具有重要意义.本研究以黄土丘陵区不同人工恢复植被为对象,以农田为参照,分析了不同人工植被恢复方式对0～100 cm剖面土壤有机碳(SOC)和全氮(TN)含量影响的差异及其影响因素.结果表明: 退耕还林还草显著提高了土壤的SOC和TN含量.退耕后,SOC和TN含量均较农田明显提高.0～100 cm土层SOC平均含量人工乔木林为农田的1.43倍,增幅最大;其次是人工灌木,为1.36倍;最后是人工草地,为1.21倍.0～100 cm土层TN平均含量人工乔木林增幅最大,是农田的1.30倍;其次是人工草地,为1.21倍;而人工灌木增幅最小,为1.13倍.与农田相比,人工恢复植被类型间SOC和TN含量及细根密度的差异在土壤剖面深度上表现出不同,人工乔木和灌木最明显,影响深度＞100 cm;草地最小,仅为60 cm.恢复植被的细根密度、C∶N和凋落物量显著高于农作物,细根密度与SOC、TN呈显著线性相关(P<0.01).细根的质和量以及凋落物量是不同人工恢复植被下SOC和TN含量差异的重要影响因素.