A model of nitrogen flows in grassland

Abstract. The model comprises three submodels, which together give an integrated picture of nitrogen pools and fluxes in grassland under grazing or cutting. The first submodel represents the interaction of the grazing animal with the sward through intake and the production of excreta: the second is concerned with the growth of the vegetative grass crop and its response to light, temperature and nitrogen; these two submodels are interfaced with a submodel of soil carbon and nitrogen pools and processes, including dead shoot and root material, dead and live soil organic matter, and three pools representing mineral nitrogen. No account is taken of water, which is assumed to be non-limiting, or the possible effects of soil pH and soil aeration. The model is used to simulate a range of management strategies as applied to stocking density and fertilizer application, examining both steady-state and non-steady-state conditions. The model highlights the long time scales associated with grassland systems, the role of the grazing animal in modifying carbon and nitrogen flows, and the importance of soil conditions to grassland productivity and fertilizer response. The productivity of grazed swards may be greater or less than that of cut swards depending on stocking density and fertilizer application, although nitrogen recovery (as calculated here) is always lower in grazed swards. The model is able to stimulate mineralization and immobilization, and place these in the context of plant processes and the grazing animal.     

草原生态系统生产力对降水格局响应的研究进展

全球变化背景下降水格局发生了显著改变,主要表现在年降水量、降水季节分配以及降水事件特征(单次降水事件雨量大小、两次降水之间的间隔时长和一年中降水事件的次数等)等几个方面,降水格局的改变将显著影响陆地生态系统功能,尤其是对生产力的影响.草原生态系统是对降水格局改变最敏感的生态系统之一,因此有必要了解草原生态系统生产力对降水格局响应的研究现状及存在的问题.本文首先综述了草原生态系统生产力对降水格局各个特征响应的国内外研究进展,进而从长期观测、控制试验、模型模拟3方面综述了目前降水格局对生产力影响的研究方法,最后从理论研究、研究方法两个方面提出了目前研究存在的问题以及未来的研究方向.本文将为预测草原生态系统如何响应未来气候变化以及在气候变化情景下进行草原生态系统管理提供重要科学依据.     

短期氮添加对祁连山亚高山草地生产力及植物多样性的影响

全球氮沉降速率的急剧增加已显著地改变了生态系统的生产力及稳定性,特别是在受N限制较严重的亚高山草地生态系统。虽然氮沉降增加对草地生产力和植物多样性影响的研究报道已经很多,但是氮素沉降的生态系效应因气候区、草地系统类型、加氮水平、氮肥类型和试验时间长短等不同而差别很大。为了评估氮沉降增加对亚高山草地植物物种多样性和生产力的影响,通过在祁连山中部亚高山草地设置不同氮添加水平（0、2、5、10、15、25 g N m^-2 a^-1和50 g N m^-2 a^-1）的短期氮沉降增加模拟试验,探讨了生产力和物种多样性对不同水平氮添加的响应。结果显示：氮添加增加了禾本科（垂穗披碱草、赖草和草地早熟禾）和莎草科（矮嵩草）的地上生产力及其在群落生产力中所占的比例,主要表现在氮添加增加了禾本科和莎草科的株高和株数,降低了其他科（鹅绒委陵菜和葛缕子）的株高和株数;与生产力相比,植物多样性对氮添加的响应较慢,总体随着氮添加量的增加呈下降趋势但未达到显著水平;植物多样性与生产力呈显著的负相关关系。研究结果表明氮添加有助于提高禾本科和莎草科的生产力,进而提高群落生产力,但其他科的植物会被逐渐替代,导致群落植物物种多样性降低。研究结果可为我国亚高山草地的持续性管理提供一定的理论基础。     

Spatial variations in aboveground net primary productivity along a climate gradient in Eurasian temperate grassland: effects of mean annual precipitation and its seasonal distribution

Concomitant changes of annual precipitation and its seasonal distribution within the context of global climate change have dramatic impacts on aboveground net primary productivity (ANPP) of grassland ecosystems. In this study, combining remote sensing products with in situ measurements of ANPP, we quantified the effects of mean annual precipitation (MAP) and precipitation seasonal distribution (PSD) on the spatial variations in ANPP along a climate gradient in Eurasian temperate grassland. Our results indicated that ANPP increased exponentially with MAP for the entire temperate grassland, but linearly for a specific grassland type, i.e. the desert steppe, typical steppe, and meadow steppe from arid to humid regions. The slope of the linear relationship appeared to be steeper in the more humid meadow steppe than that in the drier typical and desert steppes. PSD also had significant effect on the spatial variations in ANPP. It explained 39.4% of the spatial ANPP for the entire grassland investigated, being comparable with the explanatory power of MAP (40.0%). On the other hand, the relative contribution of PSD and MAP is grassland type specific. MAP exhibited a much stronger explanatory power than PSD for the desert steppe and the meadow steppe at the dry and wet end, respectively. However, PSD was the dominant factor affecting the spatial variation in ANPP for the median typical steppe. Our results imply that altered pattern of PSD due to climate change may be as important as the total amount in terms of effects on ANPP in Eurasian temperate grassland.     

氮水添加对放牧背景下荒漠草原生产力的影响

水分与氮素作为干旱和半干旱草原生产力的共同限制性因子在退化草原的生态快速修复过程中备受关注。以不同放牧强度背景下的短花针茅荒漠草原为研究对象,开展围封模拟放牧利用实验,同时添加氮素和水分。通过分析历史放牧强度与年份对生产力的影响,以及添加氮素和水分对不同功能群植物生物量的作用,探讨放牧强度对短花针茅草原生产力的内在作用机制,以及如何实现荒漠草原资源合理开发和可持续利用。研究结果显示,降雨量与放牧强度决定着短花针茅草原的植物群落结构。氮素和水分添加可分别提升11%-29%和12%-32%的群落地上生物量,且二者存在显著的交互作用。不同功能群植物的地上生物量对氮素与水分添加的响应存在差异,多年生丛生禾草对氮素和水分添加响应最敏感。氮素与水分添加可显著提高多年生丛生禾草的地上生物量,但与自然降水量相关。氮素添加对地上生物量的影响在正常降雨和稍旱年份作用显著,而水分添加在干旱年份作用显著。在正常降雨年份,以半灌木植物为优势种的轻度放牧背景以添加水分对提升生产力最宜,以多年生丛生禾草和半灌木为共优种的中度放牧背景和以多年生丛生禾草为优势种的重度放牧以同时添加水分和氮素对提升生产力最为宜;在干旱年份不同放牧强度背景下均以同时添加水分和氮素对提升生产力最为宜。我们的结果表明了养分与资源的改善有利于退化短花针茅草原的快速恢复和可持续生产。     

Regional grassland productivity responses to precipitation during multiyear above‐ and below‐average rainfall periods

There is considerable uncertainty in the magnitude and direction of changes in precipitation associated with climate change, and ecosystem responses are also uncertain. Multiyear periods of above‐ and below‐average rainfall may foretell consequences of changes in rainfall regime. We compiled long‐term aboveground net primary productivity (ANPP) and precipitation (PPT) data for eight North American grasslands, and quantified relationships between ANPP and PPT at each site, and in 1–3 year periods of above‐ and below‐average rainfall for mesic, semiarid cool, and semiarid warm grassland types. Our objective was to improve understanding of ANPP dynamics associated with changing climatic conditions by contrasting PPT–ANPP relationships in above‐ and below‐average PPT years to those that occurred during sequences of multiple above‐ and below‐average years. We found differences in PPT–ANPP relationships in above‐ and below‐average years compared to long‐term site averages, and variation in ANPP not explained by PPT totals that likely are attributed to legacy effects. The correlation between ANPP and current‐ and prior‐year conditions changed from year to year throughout multiyear periods, with some legacy effects declining, and new responses emerging. Thus, ANPP in a given year was influenced by sequences of conditions that varied across grassland types and climates. Most importantly, the influence of prior‐year ANPP often increased with the length of multiyear periods, whereas the influence of the amount of current‐year PPT declined. Although the mechanisms by which a directional change in the frequency of above‐ and below‐average years imposes a persistent change in grassland ANPP require further investigation, our results emphasize the importance of legacy effects on productivity for sequences of above‐ vs. below‐average years, and illustrate the utility of long‐term data to examine these patterns.     

降水变化与氮添加对晋北盐碱化草地土壤净氮矿化的影响

于2018年在晋北典型农牧交错带盐碱化草地设置增减降水和氮添加处理试验,2019年生长季(5-9月)采用原位顶盖PVC埋管法测定不同水、氮处理下土壤净氮矿化速率,研究盐碱化草地土壤净氮矿化速率对降水格局变化和氮沉降的响应.结果表明:土壤净氮矿化速率表现出明显的季节动态.单独增减降水(±50％)和氮添加(10g·m-2·...     

干旱胁迫-复水与氮肥耦合对小粒咖啡生长和水氮生产力的影响

以小粒咖啡(卡蒂姆P7963)为材料,研究连续2.5年不同施氮水平下周期性干旱胁迫后复水对小粒咖啡生长、产量、叶片光合特性和水氮生产力的影响.设灌水(周期性干旱胁迫后复水)和施氮2因素,4个灌水模式分别为充分灌水(I_F-F:100%ET₀+100%ET₀,ET₀为参考作物腾发量)、轻度干旱胁迫-复水(I_L-F:80%ET₀+100%ET₀)、中度干旱胁迫-复水(I_M-F:60%ET₀+100%ET₀)和重度干旱胁迫-复水(I_S-F:40%ET₀+100%ET₀),3个施氮水平分别为高氮(N_H:每次750 kg N·hm^-2)、中氮(N_M:每次500 kg N·hm^-2)和低氮(N_L:每次250 kg N·hm^-2),分4次等量施用.结果表明: 小粒咖啡株高、茎粗、产量、水氮生产力受灌水和施氮影响显著,株高和茎粗与日序数呈S型曲线关系,干旱胁迫时小粒咖啡叶片光合作用显著下降,复水后大部分光合作用指标能不同程度恢复.与I_F-F相比,I_L-F干豆产量增加6.9%,而I_M-F和I_S-F干豆产量分别减少15.2%和38.5%;I_L-F和I_M-F水分利用效率分别增加18.8%和6.0%,而I_S-F水分利用效率减少12.1%;I_L-F氮肥偏生产力增加6.1%,而I_M-F和I_S-F氮肥偏生产力分别减少14.0%和36.0%.与N_H相比,N_M干豆产量和水分利用效率分别增加20.9%和19.3%,而N_L分别减少42.4%和41.9%;N_M和N_L氮肥偏生产力分别增加81.4%和72.9%.与I_F-FN_H相比,I_L-FN_M干豆产量、水分利用效率和氮肥偏生产力分别增加37.6%、52.9%和106.4%.回归分析表明,灌水量为318 mm、施氮量为583 kg·hm^-2时,干豆产量(2362 kg·hm^-2)最大;灌水量为295 mm、施氮量为584 kg·hm^-2时,水分利用效率(0.78 kg·m^-3)最大,即产量和水分利用效率同时达到最大值时最接近I_L-FN_M水氮组合.因此,I_L-FN_M为小粒咖啡最佳的水氮组合模式.     

放牧和模拟增温对藏北高寒草地植物群落特征及生产力的影响

气候变化和放牧活动对草地植物物种多样性和生产力具有重要影响。为探索藏北高寒草地植物物种多样性和生产力对增温、放牧及其交互作用的响应, 于2011年在藏北高原开始建立增温实验平台, 2016年起增设放牧、增温+放牧实验, 连续2年(2016-2017年)观测了植物群落特征、群落组成、生产力和物种多样性。结果表明, 增温和放牧对高寒草地植物高度和净初级生产力具有显著交互作用。在放牧条件下, 增温对植物高度无显著影响; 但在不放牧条件下, 增温却显著增加了植物高度。在放牧条件下, 增温对净初级生产力的影响存在年际差异, 2016年增温对生产力无显著影响, 2017年增温显著降低了植物净初级生产力; 但在不放牧条件下, 增温对植物净初级生产力无显著影响。增温和放牧对高寒草地植物物种丰富度、盖度、重要值及多样性均无显著交互作用。植物盖度在增温和放牧条件下显著降低, 杂类草物种比例显著增加, 但物种多样性均无显著变化。研究表明, 增温和放牧显著改变高寒草地群落结构。未来气候变化条件下, 放牧活动加剧有可能导致高寒草地生产力降低。     

Small-scale pattern in grassland communities in the Serengeti National Park,Tanzania

Small-scale community gradients and patterns in four grassland communities in the Serengeti National Park, Tanzania, were analyzed by detrended correspondence analysis (DCA) and percentage distance ordination (PD). The main goals of the study were (1) to describe the vegetation patterns in the communities, (2) to identify the potential underlying causes of the patterns, and (3) to rank the communities by their relative levels of patchiness. Species cover in a small number (16–32) of discontinuous plots was estimated and soils were collected. DCA successfully identified community gradients and phases of discrete mosaics, and PD identified the relative heterogeneity of the sites. Results suggested that the two most arid sites were composed of weakly or strongly differentiated vegetational mosaics and the two most mesic sites were essentially homogeneous. Termite modification of water infiltration properties of the soils and clonal growth of dominant species were identified as two factors possibly causing the mosaic vegetation patterns.     

Relationship between road dust and ungulate density in Serengeti National Park,Tanzania

We measured the responses of ungulate grazers to roads by recording the density of dust on grasses and compared distance sampling of ungulates in Serengeti National Park (SNP), Tanzania. Data were collected on the east and west side of the Ngorongoro–Seronera main gravel road to test if road traffic and dust were important factors determining distribution patterns amongst grazers. Results indicate that dust increased progressively with traffic speed and volume during dry season. More dust was intercepted on the west than on the east side of the road mainly because of wind effects. Dust deposition (measured as density g grass g^?1 dust) was higher on short grasses than on long grasses during the dry and late‐dry seasons than during the wet season, when paired perpendicular distances up to 300 m were compared. Mean number of observed grazer species indicated that most fed further from the west side of the road than from the east perhaps to minimize higher density of dust commonly spread on foliage up to 200 m away from the road. Despite that most grazers avoided road side grass shoulders, supporting the ‘dust aversion hypothesis’, the test predictions from the ‘road disturbance’ and the ‘road attraction’ hypotheses did not support the responsive behaviours of grazers toward roads.     

Asymmetric responses of primary productivity to precipitation extremes: A synthesis of grassland precipitation manipulation experiments

Climatic changes are altering Earth's hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta‐analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.     

氮素添加对科尔沁沙质草地物种多样性和生产力的影响

物种多样性和生产力是生态系统结构和功能的重要指标.以科尔沁沙质草地为对象,通过对其进行不同梯度的氮素添加处理,研究氮素添加对沙质草地生态系统物种组成、物种多样性和生产力的影响.结果表明：氮素添加改变了群落物种组成和群落中的优势种,使植被的高度和盖度增加,植被的透光率减小;随着氮素水平的增加,群落中物种丰富度减小,物种多样性降低;不同水平的氮素添加均显著增加了(P<0.01)群落地上生物量;物种丰富度与植被透光率呈线性正相关(P<0.01),与植被盖度呈线性负相关关系(P<0.01),说明长期的氮沉降与人为氮素输入将影响沙地生态系统的物种组成、物种多样性以及生产力.     

基于BIOLOG技术分析氮沉降和降水对土壤微生物功能多样性的影响

土壤微生物是有机物分解和养分循环的主要介质,因此在维持土壤的功能多样性和持续性方面发挥着关键作用。气候变化驱动因素会影响土壤微生物的生理活动,引起其群落结构和功能多样性的改变,并对生物地球化学循环和气候―生态系统反馈产生连锁效应,其中氮沉降和降水是全球气候变化的研究热点。土壤氮(N)的有效性有可能通过改变微生物的群落组成以调节微生物对降水变化的响应,但目前关于N沉降和降水及其交互作用对土壤微生物群落功能多样性的影响机制仍不清楚。为了准确预测未来气候条件下生态系统的功能状况,需要更好地了解土壤微生物对环境变化的响应。基于BIOLOG技术综述了氮沉降和降水变化及其交互作用对土壤微生物功能多样性影响的相关研究进展,可以为进一步研究全球气候变化背景下地下生态学的发展提供参考。另外,分析阐述了当前工作中存在的一些主要瓶颈,并对未来的研究热点进行了探讨和展望。     

牧鸡密度与取样时间对沙质草地土壤氮素有效性的影响

以科尔沁沙质草地为对象,设置5个牧鸡密度D1(10羽·200 m-2)、D2(10羽·400 m-2)、D3(10羽·600 m-2)、D4(10羽·800 m-2)、CK(0羽·200 m-2),研究牧鸡密度与取样时间对土壤无机氮、氮矿化与硝化、微生物生物量等的影响.结果表明:与对照相比,高牧鸡密度(D1、D2)显著增加了8月、10月土壤潜在净氮矿化、硝化速率,而低牧鸡密度(D3、D4)对土壤潜在净氮矿化、硝化速率的影响均不显著;牧鸡显著提高了土壤硝态氮含量、土壤潜在净氮矿化、硝化速率,对土壤铵态氮含量影响不显著;土壤无机氮含量、潜在净氮矿化、硝化速率对取样时间的响应均达到显著水平,对牧鸡密度与取样时间交互作用的响应不显著.牧鸡对土壤微生物生物量碳、氮及微生物生物量碳/氮的影响均不显著.研究认为,牧鸡可以增加土壤的供氮能力,且随牧鸡密度的增加呈上升趋势,最适牧鸡密度为250 ～500羽·hm-2.     

增加降水对荒漠短命植物当年牧草生长及群落结构的影响

通过模拟降水增加试验研究了古尔班通古特荒漠常见短命植物盖度、密度及产量对水分输入量增加的响应.结果表明:尖喙牻牛儿苗、弯果胡卢巴、角果藜、琉苞菊、条叶庭芥及囊果苔草6种牧草生物量总和占总生物量的60%以上,是荒漠草原的优势牧草植物;供水量增加显著促进了角果藜、尖喙牻牛儿苗、弯果胡卢巴、条叶庭芥和琉苞菊5种牧草植物的个体生长;在自然降水的基础上,增加40mm和80mm供水使这5种植物生物产量比对照增加86%～230%.供水增加后,6种重要牧草对整个群落的生物贡献比例发生很大变化,只有角果藜和尖喙牻牛儿苗的贡献比例规律性增加,分别比对照增加了58%和11%,表明2种植物对供水增加的环境变化适应能力强;供水量增加促进了弯果胡芦巴、尖喙牻牛儿苗和角果藜3种植物N、P和K养分的吸收量;与此同时,供水量增加也显著提高了荒漠短命植物的密度和盖度.在降水增加的背景下,古尔班通古特沙漠南缘荒漠短命植物群落结构发生改变,植被盖度和群落净初级生产力提高,这对于防风固沙、稳定沙面,增加荒漠草原载畜能力有着积极的意义.     

降水时间对内蒙古温带草原地上净初级生产力的影响

全球气候变化下降水时间的改变将深刻影响草原生态系统地上净初级生产力(ANPP),而草原生态系统ANPP是区域碳循环的重要过程.利用1998-2007年的SPOT-VEG NDVI数据并结合111个样点的ANPP地面样方调查数据,获得了内蒙古温带草原1998-2007年的ANPP区域数据,依此分析了中国内蒙古温带草原以及区域内的3种植被类型(荒漠草原、典型草原、草甸草原)降水时间对ANPP的影响.研究结果表明,对于整个内蒙古温带草原来说,一个水分年内(从上一年9月份到当年地上生物量达最大值时的8月份)影响ANPP较为重要的降水月份为2-7月份,其中,5-7月份降水尤为重要.具体到每个月降水的影响,研究发现,7月份降水最重要,而仍处于生长季的8月份降水相对于其他生长季降水作用最小;影响不同草地类型最重要的降水时期存在一定差异,对荒漠草原和典型草原地区来说,ANPP达最大值前3个月(5-7月份)的生长季降水最重要,而8月份降水影响较小,而草甸草原地区8月份和非生长季的3、4月份降水最重要,但各个降水时期降水对ANPP的影响都较荒漠草原和典型草原小,大部分地区降水对ANPP的影响不显著.     

降水和氮沉降增加对草地土壤酶活性的影响

为探究降水和氮沉降增加对草地生态系统土壤酶活性的影响,于2014年生长季在内蒙古温带典型羊草草原开展了野外原位控制实验。试验共设置降水(对照,W0,自然降水;W15,增加15%的年均降水量)、施氮(对照,CK,0 kg N hm~(-2)a~(-1);低氮,LN,25 kg N hm~(-2)a~(-1);中氮,MN,50 kg N hm~(-2)a~(-1);高氮,HN,100 kg N hm~(-2)a~(-1))及其交互作用等8个不同的处理水平来模拟降水和氮沉降增加的全球变化情景,分别定量探讨了不同水、氮添加条件下草地表层土壤中与氮循环相关的蛋白酶,脲酶,硝酸还原酶,亚硝酸还原酶活性的月动态变化及其与土壤理化性质之间的相关性。研究结果表明:在自然降水条件下,不同施氮水平蛋白酶、脲酶和硝酸还原酶活性无显著差异,亚硝酸还原酶活性相比于对照显著降低;在增加降水条件下,不同施氮水平对蛋白酶和硝酸还原酶活性未产生显著性影响,高氮水平显著降低脲酶和亚硝酸还原酶活性。不同施氮水平是否添加降水对亚硝酸还原酶活性无影响,而增添降水使低氮处理的蛋白酶活性和中、高氮处理水平的硝酸还原酶活性增加、高氮处理的脲酶活性降低。降水在影响蛋白酶和硝酸还原酶活性方面具有主效应,氮沉降在影响亚硝酸还原酶活性方面具有主效应,而降水和施氮处理未表现出明显地交互作用。土壤亚硝酸还原酶活性与土壤碳氮比和NH~+_4-N含量极显著正相关,与NO-3-N含量显著正相关。