不同降水条件下荒漠草原植物的养分含量及回收对增温和氮素添加的响应

为了探讨荒漠草原植物养分回收特征对长期增温和氮素添加的响应以及自然降水变异对其的调控作用,该研究依托实施12年的模拟增温和氮素添加实验平台,在相对多雨的2016年(超过长期均值52%)和相对少雨的2017年(低于长期均值16%),以常见C_3植物银灰旋花(Convolvulus ammannii)和C_4植物木地肤(Kochia prostrata)为研究对象,测定分析绿叶和枯叶的氮磷含量及回收效率。结果表明:(1)在相对多雨年(2016年),增温使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别增加了14.32%、25.45%、17.97%和46.47%,氮、磷回收效率分别显著减小了9.41%和16.99%(P0.05);氮素添加使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别提高了17.32%、25.62%、20.21%和51.41%,而氮、磷回收效率显著降低了9.33%和18.89%(P0.05);增温+氮素添加共同处理显著增加了植物氮磷含量、降低了氮磷回收效率。(2)在相对少雨年(2017年),增温、氮素添加、增温+氮素添加处理对植物叶片氮磷含量、回收效率均无显著影响。(3)叶片氮磷含量在物种间差异极显著(P0.000 1),而氮磷回收效率在物种间无显著差异。(4)回归分析表明,植物叶片氮磷含量随着土壤无机氮、有效磷及含水量的增加而增加,植物氮磷回收效率则随着土壤养分和水分的可利用性的增加而降低。研究认为,荒漠草原植物养分回收对全球变化的响应受自然降水变异的调控。     

中国北部半干旱区乔木、灌木和草本3种不同生活型植物的氮素回收特征

在位于中国北部半干旱区的多伦县选择 3种不同生活型的植物 ,每种生活型各选取 3种有代表性的植物 :3种落叶乔木白桦 Betula platyphylla 、山定子 Malus baccata 、山杏 Prunus armeniaca ,3种落叶灌木黄柳 Sal-ix flavida 、山刺玫 Rosa davurica 、羊柴 H edysarum laeve 和 3种多年生草本地榆 Sanguisorba officinalis 、菊叶委陵菜 Potentilla tanacetifolia 、叉分蓼 Polygonum divaricatum ,对它们的氮素回收特征进行了研究 .结果表明 :乔木、灌木及草本 3种不同生活型植物的氮素回收效率分别为 4 3.4 2 %、5 8.84 %、73.76 % ,氮素回收水平分别为 12 .4 m g· g- 1、10 .7mg· g- 1和 8.2 m g· g- 1 .两种深根系生活型植物 乔木、灌木 的枯叶具有较高的氮素浓度 即回收水平低 ,并且成熟绿叶与枯叶中的氮素浓度差异较小 即回收效率低 ,与之相比 ,浅根系的多年生草本植物的枯叶具有较低的氮素浓度 即回收水平高 ,并且成熟绿叶与枯叶中的氮素浓度差异较大 即回收效率高 .不同生活型植物氮素回收能力的差异说明了不同生活型植物对生境具有不同的氮素适应策略 ,另一方面 ,氮素回收效率和氮素回收水平可作为衡量植物氮素回收能力的两个重要参数 .     

氮素和水分添加对贝加尔针茅草原植物多样性及生物量的影响

为了解草原植物群落物种多样性和植物地上生物量对氮沉降增加和降水变化的响应,在内蒙古贝加尔针茅(Stipa baicalensis)草原,分别设置对照(N0)、1.5 g/m2(N15)、3.0 g/m2(N30)、5.0 g/m2(N50)、10.0 g/m2(N100)、15.0 g/m2(N150)、20.0g/m2(N200)和30.0 g/m2(N300)(不包括大气沉降的氮量)8个氮素(NH4NO3)添加梯度和模拟夏季增加降水100mm的水分添加交互试验,研究氮素和水分添加对草原群落植物物种多样性和几种常见植物地上生物量的影响。结果表明:(1)氮素和水分的添加降低了草原群落植物物种多样性,且氮素和水分有显著的互作效应。在水分添加的条件下,随着施氮水平的增加,群落植物物种多样性减小;在无水分添加的条件下,随着施氮水平的增加,群落植物物种多样性呈先增加后减小的"单峰"变化趋势。(2)不同植物对氮素和水分添加的响应不同,随着施氮水平的增加,羊草地上生物量显著增加;贝加尔针茅、羽茅、糙隐子草、寸草苔和冷蒿先增加后减少,呈单峰曲线;星毛委陵菜、牧马豆、扁蓄豆和线叶菊地上生物量则逐渐减少。而且氮素和水分对贝加尔针茅、羽茅、扁蓄豆地上生物量有显著的交互作用。     

亚热带湿地松叶片多元素化学计量与养分回收对氮添加的短期响应

在我国南方亚热带湿地松人工林设置了3个水平的野外氮添加控制试验(0、40、120 kg N·hm^-2·a^-1),于2014和2015年生长季高峰期(7月底)和末期(10月底)采集湿地松成熟绿叶和落叶,分析外源氮添加对湿地松叶片碳(C)、氮(N)、磷(P)、钾(K)、钙(Ca)、镁(Mg)、铝(Al)、铁(Fe)、锰(Mn)9种元素浓度及其养分回收的影响.结果表明: N添加显著增加了湿地松绿叶中N、Al、Mn浓度,降低了P和2014年的Ca浓度,而对C、K、Mg、Fe 浓度无显著影响.N添加显著提高了绿叶N/P,且该比值及绿叶养分浓度(N、P、Mn)对N添加的响应依赖于N的剂量(高N条件下响应更强).N添加显著降低了2015年N的回收效率,提高了2014年K的回收效率.相比于养分回收效率,回收能力对增加的可利用氮响应更强.N添加显著降低了N的回收能力,提高了P、K的回收能力,降低了枯叶中的Fe浓度,而对枯叶中Ca、Mg、Al、Mn浓度无显著影响.这表明,N添加对叶片化学计量的影响因不同元素而异,植物会通过调整自身的养分内循环(养分回收)来应对环境变化.N添加提高了绿叶N/P和K/P,说明氮添加条件下植物生长可能由N、P共同限制转变为P限制.氮添加增加了绿叶中Al、Mn浓度,表明N添加下湿地松面临潜在的金属离子毒性风险升高.     

内蒙古典型草原羊草群落氮素去向的示踪研究

 在中国科学院内蒙古草原生态系统定位研究站的羊草样地,采用15N同位素示踪技术研究了羊草（Leymus chinensis）群落标记氮素的去向。结果表明：在我国典型草原羊草群落,植物对标记氮素的回收率为31.61%,氮素添加显著影响植物对标记氮素的回收,随着氮素添加量的增加,地上和地下植物器官对标记氮素的回收量均显著提高。标记氮素被凋落物的回收率为2.92%,地下凋落物的回收率显著高于地上凋落物。标记氮素的土壤存留率为36.16%,主要分布在地表0～40 cm的土层范围内;各土层存留的标记氮素量均随着氮素添加量的增加而显著提高。标记氮素的当季损失率为21.77%～43.38%。风险/收益比分析表明,在该试验条件下,添加5.25 g N•m-2与28 g N•m-2的处理风险大于收益,添加17.5 g N•m-2的处理风险最低,收益最高,在草原生态系统的管理中可供参考。     

模拟增温和氮素添加对荒漠草原生态系统碳交换的影响

气候变暖和大气氮沉降是全球变化的重要驱动因子。在草地生态系统中,气温升高和大气氮沉降都会改变草地固碳(C)状况,然而温度增加和大气氮沉降是如何影响生态系统碳交换目前还不明确。本研究旨在研究增温和氮素添加对荒漠草原碳交换的影响。在短花针茅荒漠草原上采用2×2因素完全随机区组的裂区设计,使用红外辐射器来模拟气候变暖并且使用添加氮肥的方法来模拟大气氮沉降,在不同处理条件下测定生态系统净碳交换(NEE)、生态系统呼吸(ER)和总生态系统生产力(GEP),分析了2013和2014年影响短花针茅荒漠草原生态系统C交换的因素,结果如下:(1)增温使土壤温度显著增加了0.70℃(P0.001),土壤湿度显著增加了7.58%(P0.001)。(2)增温、氮素添加及其交互作用显著增加了GEP和ER(P0.05),而对于NEE没有显著影响(P0.05)。(3)2013年GEP在8月初达到峰值,ER在8月末9月初达到峰值,NEE随着GEP和ER的变化而波动;2014年GEP、ER和NEE均在8月末9月初达到峰值。(4)ER和GEP随着大气温度升高和降水增加而增大,土壤温度和土壤湿度也是影响生态系统C交换的重要因素。     

羌塘高原降水梯度带紫花针茅叶片氮回收特征及影响因素

植物回收衰老叶片的氮是植物重要的养分保持和环境适应机制,在寒旱贫瘠的生境更是如此。为了理解降水梯度上植物对高寒贫瘠环境的养分适应特征,研究了羌塘高寒草原优势物种紫花针茅叶片氮回收策略及其与环境因子的关系。结果表明,降水梯度带上紫花针茅叶片具有较高的叶氮水平和氮回收能力。生长季盛期紫花针茅绿叶平均氮含量为(23.87±3.92)g/kg,高于中国草地平均水平(20.9 g/kg)及全球平均值(20.1 g/kg);绿叶氮含量与年降水量(MAP)呈显著负相关,干旱端(西部)绿叶中氮含量明显高于湿润端(东部)。枯叶养分回收后的氮水平(NRP)很低,平均为(6.76±1.42)g/kg,叶片平均氮回收效率(NRE)为(71.25±6.46)%,明显高于中国温带草原和全球的平均水平(46.9%—58.5%)。枯叶中氮回收水平对叶片氮回收效率起决定作用,是维持高养分回收效率的物质基础。NRE与MAP、土壤全氮(TN)和土壤无机氮呈显著负相关;NRP与TN相关性不显著,但与土壤无机氮显著负相关。尽管NRE与NRP呈显著负相关,但二者与绿叶氮含量均没有显著相关性。年均气温、海拔对NRE和NRP影响均不显著。因此,紫花针茅叶片极高的NRE和低NRP反映了它对极端干旱贫瘠环境的养分保持能力,通过内部氮循环来降低养分流失。土壤氮的有效性是影响紫花针茅叶片氮回收能力的关键因子,降水通过影响土壤氮的有效性以及绿叶中氮含量间接影响紫花针茅叶片氮回收效率。     

增温和氮素添加对内蒙古荒漠草原植物生殖物候的影响

植物物候是响应气候变化的敏感生物指示性状,在气候变暖并伴随着大气氮沉降增加的背景下,物候变化是气候变迁导致群落结构和生态系统功能变化的重要机制。本研究在以短花针茅(Stipa breviflora)为建群种的内蒙古荒漠草原设置了增温、氮素添加及其交互作用的实验平台,对短花针茅、木地肤(Kochia prostrata)、银灰旋花(Convolvulus ammannii)和细叶韭(Allium tenuissimum)4种多年生优势植物的开花、结实等物候期和生殖生长持续时间进行连续3个生长季(2008—2010年)的观察,植物物候观测采用定株标记,并对各植株上所有物候状态进行打分并以其非加权平均值作为植株整体物候分值,全年观测结束后将植株的物候分值随时间的变化利用Richards方程进行拟合,最终根据回归方程计算植物各物候期。结果表明:1)在3个生长季中,增温使4个植物种的平均开花、结实时间分别显著地提前了2.28和1.88 d·a-1;2)氮素添加对这些植物种的开花和结实时间都没有显著性影响;3)同时进行增温和氮素添加处理使植物的平均开花、结实时间分别提前1.75和1.53 d·a-1,但2种处理之间没有交互作用;4)增温使这些植物的平均生殖生长持续时间显著延长0.84 d·a-1,使短花针茅和木地肤生殖生长持续时间分别延长了1.20和3.25d·a-1。氮素添加使细叶葱的平均生殖持续时间显著缩短了1.52 d·a-1。这些结果揭示了气温升高和氮沉降影响下荒漠草原优势植物生殖物候的变化,将为分析不同种植物对各类资源需求规律的变化提供直接的实验证据,对于进一步预测全球变化背景下种间竞争格局和群落结构的演变趋势具有重要参考价值。     

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。     

四翅滨藜叶片C、N、P生态化学计量特征对N添加的响应

以宁夏平罗西大滩四翅滨藜人工林为研究对象,通过设置N添加的野外实验,研究四翅滨藜叶片C、N、P化学计量比的季节动态及其对N添加的响应特征。结果显示:(1)四翅滨藜叶片C、N、P化学计量比在生长季初期和末期较高,在生长季旺期(8～9月)较低。(2)N添加提高了绿叶N浓度和N∶P比,降低了绿叶C∶N、N回收度(NRP)和P回收度(PRP),对其他指标的影响无明显的规律性。(3)N回收效率(NRE)和NRP均与枯叶C∶N比显著正相关;P回收效率(PRE)与绿叶P浓度显著正相关,与枯叶P浓度显著负相关;PRP分别与绿叶P浓度和枯叶C、N、P化学计量比显著正相关,与枯叶C浓度显著负相关。研究表明,N添加促进了四翅滨藜绿叶N摄取,降低了叶片从枯叶中回收N和P的能力,改善了枯叶N分解质量;未来大气N沉降增加会改变干旱半干旱区植物N吸收、分配和回收等策略,促进枯叶中N的释放速率,直接影响N循环,进而间接影响到植被-土壤系统C和P的循环过程。     

Nitrogen economy of alpine plants on the north Tibetan Plateau: Nitrogen conservation by resorption rather than open sources through biological symbiotic fixation

Nitrogen (N) is one of the most important factors limiting plant productivity, and N fixation by legume species is an important source of N input into ecosystems. Meanwhile, N resorption from senescent plant tissues conserves nutrients taken up in the current season, which may alleviate ecosystem N limitation. N fixation was assessed by the ¹⁵N dilution technique in four types of alpine grasslands along the precipitation and soil nutrient gradients. The N resorption efficiency (NRE) was also measured in these alpine grasslands. The aboveground biomass in the alpine meadow was 4–6 times higher than in the alpine meadow steppe, alpine steppe, and alpine desert steppe. However, the proportion of legume species to community biomass in the alpine steppe and the alpine desert steppe was significantly higher than the proportion in the alpine meadow. N fixation by the legume plants in the alpine meadow was 0.236 g N/m², which was significantly higher than N fixation in other alpine grasslands (0.041 to 0.089 g N/m²). The NRE in the alpine meadows was lower than in the other three alpine grasslands. Both the aboveground biomass and N fixation of the legume plants showed decreasing trends with the decline of precipitation and soil N gradients from east to west, while the NRE of alpine plants showed increasing trends along the gradients, which indicates that alpine plants enhance the NRE to adapt to the increasing droughts and nutrient‐poor environments. The opposite trends of N fixation and NRE along the precipitation and soil nutrient gradients indicate that alpine plants adapt to precipitation and soil nutrient limitation by promoting NRE (conservative nutrient use by alpine plants) rather than biological N fixation (open sources by legume plants) on the north Tibetan Plateau.     

Resorption efficiency of leaf nutrients in woody plants on Mt. Dongling of Beijing,North China

Aims To explore resorption efficiency of nitrogen (NRE) and phosphorus (PRE) of woody plants in relation to soil nutrient availability, climate and evolutionary history, in North China.Methods We measured concentrations of nitrogen ([N]) and phosphorus ([P]) in both full expanded mature green and senescent leaves of the same individuals for 88 woody species from 10 sites of Mt. Dongling, Beijing, China. We built a phylogenetic tree for all these species and compared NRE and PRE among life forms (trees, shrubs and woody lianas) and between functional groups (N-fixers and non-N-fixers). We then explored patterns of NRE and PRE along gradients of mean annual temperature (MAT), soil inorganic N and available P, and phylogeny using a general linear model.Important findings Mass-based NRE (NRE m) and PRE (PRE m) averaged 57.4 and 61.4%, respectively, with no significant difference among life forms or functional groups. Neither NRE m nor PRE m exhibited significant phylogenetic signals, indicating that NRE m and PRE m were not phylogenetically conserved. NRE m was not related to [N] in green leaves; PRE m was positively correlated with [P] in green leaves; however, this relationship disappeared for different groups. NRE m decreased with [N] in senescent leaves, PRE m decreased with [P] in senescent leaves, for all species combined and for trees and shrubs. NRE m decreased with soil inorganic N for all species and for shrubs; PRE m did not exhibit a significant trend with soil available P for all species or for different plant groups. Neither NRE m nor PRE m was significantly related to MAT for overall species and for species of different groups.     

江西阳际峰30种阔叶树叶片氮磷含量及再吸收效率

养分再吸收是植物养分利用的重要策略,体现了植物对养分留存、利用和适应环境的能力。为研究亚热带不同生活型(常绿与落叶)阔叶树养分含量与养分再吸收的关系,以江西阳际峰国家级自然保护区内30种阔叶树为研究对象,测定成熟和衰老叶片氮(N)和磷(P)含量,分析常绿和落叶树种叶片N和P含量及其再吸收效率差异,揭示阔叶树种叶片养分再吸收效率对植物生活型的响应。结果表明: 落叶树种成熟叶片N和P含量显著高于常绿树种,衰老叶片P含量显著高于常绿树种,而两者衰老叶N含量差异不显著;30种阔叶林木叶片的氮再吸收效率(NRE)与磷再吸收效率(PRE)平均值分别为49.6%和50.9%,两种生活型树种间叶片的NRE与PRE无显著差异;落叶和常绿树种叶片的NRE均与衰老叶N含量呈显著负相关,PRE则与衰老叶P含量呈显著负相关,且这种关系在不同生活型之间差异不显著;总物种的PRE-NRE异速生长指数为1.18。江西阳际峰30种不同生活型阔叶树的养分再吸收效率会影响衰老叶片的养分状况,且相较于N,植物偏好从衰老叶中再吸收P。     

Plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming

Plant nitrogen (N) relationship has the potential to regulate plant and ecosystem responses strongly to global warming but has not been carefully examined under warmed environments. This study was conducted to examine responses of plant N relationship (i.e. leaf N concentration, N use efficiency, and plant N content in this study) to a 4‐year experimental warming in a tallgrass prairie in the central Great Plains in USA. We measured mass‐based N and carbon (C) concentrations of stem, green, and senescent leaves, and calculated N resorption efficiency, N use efficiency, plant N content, and C : N ratios of five dominant species (two C₄ grasses, one C₃ grass, and two C₃ forbs). The results showed that warming decreased N concentration of both green and senescent leaves, and N resorption efficiency for all species. N use efficiencies and C : N ratios were accordingly higher under warming than control. Total plant N content increased under warming because of warming‐induced increases in biomass production that are larger than the warming‐induced decreases in tissue N concentration. The increases in N contents in both green and senescent plant tissues suggest that warming enhanced both plant N uptake and return through litterfall in the tallgrass ecosystem. Our results also suggest that the increased N use efficiency in C₄ grasses is a primary mechanism leading to increased biomass production under warming in the grassland ecosystem.     

Nitrogen resorption from needles of Pinus thunbergii Parl. growing along a topographic gradient of soil nutrient availability

To examine the relative importance of nutrient resorption in increasing the nutrient-use efficiency of Pinus thunbergii Parl., we investigated the nitrogen contents of green and senescent needles of P. thunbergii trees growing at five positions along a slope (LS, lower slope; TR, transitional site; MS, middle slope; US, upper slope; RG, ridge) and found that soil nitrogen availability tended to decrease upslope. Nitrogen concentration in green and senescent needles decreased upslope. Nitrogen resorption efficiencies (percentage change in nitrogen content between green and senescent needles) increased upslope from 43 to 77% with decreasing soil nitrogen availability. Nitrogen resorption efficiency was related to green needle dry mass per unit length, but there was no clear correlation between nitrogen resorption efficiency and nitrogen content in green needles. We concluded that the increase in nitrogen resorption efficiency of P. thunbergii enhanced the nitrogen-use efficiency as a response to the low nitrogen availability.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

亚热带幼林树木功能性状与叶片氮磷重吸收率的关系

本研究以亚热带29种3年生人工纯林为对象,研究了29个树种功能性状与氮磷重吸收效率的关系。结果表明: 29种幼林平均氮、磷重吸收效率分别为50.5%和57.3%。22种丛枝菌根树种的氮重吸收效率平均为52.7%,显著高于7种外生菌根树种(45.1%)。29个树种的细根组织密度与氮重吸收效率呈显著正相关,7种外生菌根树种细根直径与磷重吸收效率呈显著正相关,22种丛枝菌根树种的功能性状对氮重吸收效率和磷重吸收效率无显著影响。在29个树种中,菌根类型、比叶面积、细根组织密度、叶厚度及叶厚度与菌根类型的相互作用共同解释氮重吸收效率变异的27%,比根长、细根碳含量、细根碳氮比、菌根类型、叶片碳含量及叶片碳含量与菌根类型的相互作用共同解释磷重吸收效率变异的35%。因此,亚热带树种根系功能性状能较好地预测了氮、磷养分重吸收效率,综合多个功能性状可以更好地揭示不同生物因子对养分重吸收的相对重要性。