内蒙古高原成熟和退化羊草草原群落物种功能特性与土壤微生物量C、N、P对氮素添加响应

氮素是大多数陆地生态系统初级生产力的主要限制因子。由于人类的工业和农业生产活动不断加剧，导致全球性氮沉降增加，使大多数生态系统氮素的可获得性增强。从而降低或消除了氮素对生态系统的限制作用，加速了生态系统生物地球化学过程，对物种多样性和生态系统结构与功能产生了显著的影响。但由于成土母质、气候条件、地形地貌、植被组成等的差异，不同生态系统类型对氮素增加的响应也不尽相同。欧洲和北美一些发达国家地区对于草地生态系统对于全球性氮沉降增加响应进行了较全面的研究，对于分布广泛的欧亚大陆草原研究相对不足。 本文研究选择对于欧亚大陆草原较具代表性的成熟羊草草原群落及该群落的退化类型为研究对象，从1999年开始，在这两类群落中选取地形相对平缓均一，植被组成一致的地段设置了施肥小区并进行持续氮素添加实验。本文研究了成熟和退化羊草草原群落物种功能特性与土壤微生物量C、N、P对氮素添加响应。 羊草群落中6种主要植物的地上生物量、种群密度、比叶面积、叶氮和叶绿素含量对于氮素添加响应以及各指标之间相关关系的分析表明：比叶面积、基于质量的叶片含氮量和叶绿素含量、叶绿素a和叶绿素b的比值等叶片水平上物种功能特性间的相互作用，共同影响和决定了种群密度和地上生物量对氮素添加的响应。羊草通过提高比叶面积、叶片叶绿素含量和含氮量、种群密度及个体生物量等多重调节功能对氮素添加做出响应。西伯利亚羽茅主要通过提高比叶面积、单位质量叶片的叶绿素含量和含氮量，以及株丛生物量，使其在群落占据优势。大针茅和冰草在提高比叶面积、叶片叶绿素含量和含氮量的调节能力相对较低，种群密度沿氮素添加梯度显著降低。黄囊苔草只能通过提高叶片叶绿素含量和含氮量对氮素添加做出响应，其叶绿素a与叶绿素b的比值沿氮素添加梯度逐渐降低，种群密度和地上生物量也显著降低。糙隐子草的叶绿素a与叶绿素b比值沿氮素添加梯度显著降低，但由于糙隐子草具有较高的SLA，且对叶绿素、叶片含氮量的调节能力较强，氮素添加处理没有对其种群密度和地上生物量产生显著的影响。上述结果支持Tilman的光资源竞争假说和Knops等的物种替代假说。 成熟和退化羊草群落土壤微生物量、土壤有机碳、全氮、全磷、速效氮、pH以及凋落物碳、氮、磷含量的测定结果表明：（1）成熟羊草群落表层土壤微生物量碳、氮、磷含量均随氮素添加量的增加而降低;退化羊草群落表层土壤微生物量碳、氮、磷含量沿氮素梯度表现出先增加而后降低的趋势;相关分析的结果显示各群落土壤微生物量碳、氮、磷均与土壤pH呈显著的正相关。（2）微生物量碳、氮、磷含量均随土层深度的增加而下将;而对照的微生物量碳、氮、磷含量则与土壤有机质含量呈显著正相关。（3）年度间降水量差异对土壤微生物量碳、氮、磷具有较大影响。综合上述研究结果，我们认为成熟羊草群落土壤微生物生长不受氮素限制，但退化群落不同;氮素添加导致的土壤酸化作用可能是两类群落表层土壤微生物量下降的主要因素，且这种影响主要集中在0-10cm的表层土壤;表层土壤微生物量碳、氮、磷对氮素添加的响应可能还受到其它因子（如生长季降水量）的影响;深层土壤微生物量较低主要是由于土壤有机质含量较低的缘故。

黄土丘陵区草地土壤微生物C、N及呼吸熵对植被恢复的响应

以野外样地调查和室内分析法研究了黄土丘陵区不同植被恢复年限下草地土壤微生物C、N及土壤呼吸熵的变化。结果表明,土壤微生物量碳明显地随着植被恢复年限的增加而增加。在恢复前23a,土壤微生物量碳在0～20cm土层年增加率为24.1%;20～40cm为104.4%。植被恢复23a后,0～20cm土层增长率为0.83%,20～40cm为0.19%。土壤微生物量N表现为在植被恢复的初期略有下降,3a后,开始出现明显增加。0～20cm土层年增长率为20.14%,20～40cm为15.11%。在植被恢复23a后,0～20cm土层的年增长率为0.14%,20～40cm变化不大。土壤微生物呼吸强度随着恢复年限的增加逐渐加强;土壤呼吸熵随植被封育时间的增加而呈对数降低趋势。土壤呼吸熵(qCO2)在反映土壤的生物质量变化时,显得更加稳定,受植物生长状况影响较小。相关分析表明,土壤微生物量和土壤微生物活性与土壤有机质、碱解氮和粘粒含量显著正相关;与土壤粉粒含量明显负相关;表层土壤pH值对其也有明显影响。草地植被自然恢复过程可增加土壤微生物活性,有利于土壤质量的提高。

开垦和放牧对内蒙古草原土壤碳库和温室气体通量的影响

本文采用野外观测和室内测定相结合的方法，研究了内蒙古草原两种主要的土地利用方式（开垦和放牧）对土壤碳库和温室气体通量的影响，结果如下： 1．内蒙古草甸草原开垦后30年后，与天然草原相比，在休闲年份0～20 cm土壤有机碳和土壤微生物量碳含量均没有显著下降，但开垦导致土壤易分解碳下降了24%。土壤易氧化碳受季节性水热因子的驱动，表现出明显的季节变化动态。因此，土壤易分解碳的较土壤碳库的其他组分对开垦更敏感，是表征土壤管理措施引起有机质变化的一个重要指标。 草原开垦后，土壤-植物系统氧化大气甲烷的能力明显提高，农田和天然草原CH4平均吸收通量分别是48.9 和 29.0 μg C m-2 h-1，开垦后增加了1.7倍。开垦没有改变CH4吸收通量 “夏季高秋季低” 的季节变化规律（由降水决定）;开垦使N2O的平均释放通量增加了47%，农田和天然草原N2O平均吸收通量分别是56.6 和 38.6 ugN m-2 h-1;开垦同时也增大了通量的变异幅度;但没有改变N2O季节变化规律，只是出现高峰的时间较天然草原推后约10天左右。 2．开垦后的农田土壤在模拟添加厩肥后，刺激了土壤微生物的呼吸代谢，使CO2的释放量增加了5-7倍。试验期间总体排放的CO2中，约60%来源于羊粪，40% 来源于土壤。两种土壤即羊草顶级草原土壤（高碳高氮）和冷蒿－小禾草退化草原土壤（低碳低氮），在CO2的释放总量和释放比例上都没有显著性差异。添加厩肥均造成两种土壤碳库的净碳损失，并且退化草原土壤（7.0%）的土壤净碳损失要大于羊草草原（2.6%）。说明与开垦后的高C土壤相比，在已经退化草原的低C土壤上施厩肥将趋向于土壤更大的净碳损失。 3．自由放牧22年后，羊草草原0～10 cm土壤有机碳、微生物量碳和易分解碳分别下降了14.1%、27.9%和22.0%;大针茅草原0～5 cm土壤有机碳和微生物量碳分别下降了27.6%和38.2%。两类草原土壤碳组分的季节变化受水热因子的驱动，大针茅草原季节波动出现高峰的时间较羊草草原迟。土壤微生物量碳在表征羊草草原和大针茅草原土壤碳素的动态变化时，要敏感于土壤总有机碳。放牧对冷蒿-小禾草草原土壤各碳素组分影响不明显。在表征放牧对冷蒿-小禾草草原土壤的影响指示上，MB-C/ Org-C和Lab-C/ Org-C要比MB-C和Lab-C更加敏感。这说明在研究放牧对草原土壤碳库影响时，不同的草原类型应使用不同的指标来表征其变化。 内蒙古羊草草原是大气CH4的汇，自由放牧增加土壤对CH4的吸收。CH4平均吸收通量增加了27%，但CH4吸收的季节变化形式没有改变;放牧使。自由放牧还增大了N2O的排放通量，将原来N2O源、汇的双重功能改变为单一的源功能;放牧使N2O平均释放通量增加了1倍;放牧显著增加了羊草草原向大气排放CO2的量（p<0.05），并且年度排放量范围也有所增大。 4．草原羊尿斑土壤的pH和NH4+浓度在施后显著升高，但土壤微生物C和N没有明显变化，尿斑N素会发生大量的流失。粪斑和厩肥斑中各有46%和27%的N素分解后转移到植物中。羊草种群斑块上粪尿斑引起CO2和N2O通量的变化，要大于星毛委陵菜种群斑块。与植被类型的影响相比，羊粪尿斑尤其是尿斑对温室气体通量的影响更大。尿斑既降低了土壤对CH4的吸收，又增加了CO2和N2O的释放，使粪尿斑上相当于CO2的净排放量比对照土壤增加了15%。 在内蒙古草原中等放牧条件下，家畜粪尿斑在放牧草地上的覆盖面积约只有2%，与未被家畜排泄物覆盖的草原土壤 相比，粪尿斑对内蒙古草原温室气体总体收支产生的影响可以忽略不计。因此内蒙古草原地区温室气体减排措施的重点，应放在家畜的食性食量对温室气体的影响以及厩肥的科学利用上。但随放牧强度的加大，家畜排泄物覆盖草地的面积将大大增加，加之放牧生态系统中家畜瘤胃代谢产生的的大量温室气体，其对草原温室气体的核算将会产生的影响也是不容忽视的。

黄土旱塬区冬小麦不同施肥处理的土壤呼吸及土壤碳动态

依据黄土旱塬区黑垆土上中国科学院长武站长期定位试验(始于1984年),于2008年3月到6月,测定了冬小麦连作系统中返青期、拔节期、抽穗期、灌浆期和收获期土壤呼吸日变化、生育期变化以及土壤可溶性有机碳(Dissolved organic C,DOC)和微生物量碳(Soil microbial biomass C,MBC),研究了施肥措施对土壤呼吸、DOC和MBC的影响以及土壤呼吸与碳组分之间的关系。研究涉及6个处理:休闲地(F)、不施肥(CK)、有机肥(M)、氮肥(N)、氮磷肥(NP)和氮磷有机肥(NPM)。结果表明,冬小麦连作系统中土壤呼吸的日变化格局呈单峰曲线,最高值出现在12:00左右(拔节期)和14:30左右(成熟期),最小值出现在0:00～3:00之间或6:00左右;冬小麦土壤呼吸速率拔节期最高,其次是灌浆后期,抽穗期最低;不同施肥条件下,各生育期土壤呼吸速率大小顺序:NPM>M>NP>N>CK>F。土壤水分亏缺是导致抽穗期和灌浆期土壤呼吸速率降低的重要原因。各施肥处理DOC含量高低顺序为灌浆期>抽穗期>成熟期>返青期>拔节期;除M,NPM处理MBC含量拔节期>灌浆期外,各施肥处理MBC含量高低顺序...

岷江上游人工针叶、阔叶林土壤可溶性有机物质特征及其对凋落物分解的响应

土壤可溶性有机物质（Dissolved organic matter，DOM）作为土壤有机质的活性组分，在陆地生态系统物质循环中扮演非常重要的角色。土壤DOM的主要成分可溶性有机碳（Dissolved organic carbon，DOC）和氮（Dissolved organic nitrogen，DON）参与C、N循环过程。为深入揭示全球C、N循环过程机制，在未开展DOC和DON的地区进行相关研究是有必要的。森林土壤（包括枯枝落叶层）DOC、DON动态及调控机理的研究是目前国际上森林生态系统C、N循环研究热点之一。本研究立足于暖温带岷江上游茂县地区人工林植被，对土壤DOC和DON的库容量，季节动态及其与其它养分之间的关系进行了系统研究，旨在了解DOC和DON在该区生态系统中的重要作用，并探讨作为DOM主要来源的叶凋落物对DOC和DON的动态影响，研究有助于更加详细地了解该区生态系统C和N循环过程。本论文主要研究结论如下： 1研究了岷江上游地区两大主要土壤类型（棕壤和褐土）不同植物群落下土壤的DOC和DON含量及特征，结果表明：DOC和DON在两种土壤类型中均有库容量存在，DOC在0-10cm和10-20cm土层的含量幅度分别111.96~159.95 mg kg-1和69.02~100.84 mg kg-1。DON在0-10cm和10-20cm土层的含量幅度分别11.88~23.08 mg kg-1和4.70~10.77 mg kg-1。游离氨基酸在0-10cm和10-20cm土层的含量幅度分别0.84~1.66 mg kg-1和0.39~0.73 mg kg-1。DOC、DON与土壤中的一些养分因子表现出了显著的相关关系，共同反映了土壤的状况和质量，在该区开展DOC和DON的系统研究是有必要的。 2 对油松与连香树林地土壤DOC、DON以及其它化学指标的季节动态进行了研究，结果表明：油松与连香树林地土壤DOC和DON的季节动态变化表现了类似的规律，DOC和DON的含量均以秋季最高。DOC和DON的季节动态变化主要受凋落物生物因素的影响，但其微生物活力的生物因素以及降雨、温度等非生物因素也是控制土壤DOC和DON含量的重要因素。土壤DON在土壤中的行为不同于矿质氮，其季节动态不同于NO3--N和NH4+-N的季节动态，在研究N循环过程中，应考虑DON的变化情况。 3 对油松与连香树林地分解层和表层土壤（0-10cm）氨基酸周转动态进行了研究，结果表明：油松林地和连香树林地均以分解层的氨基酸含量高于矿质表层土壤的含量。每个取样时期，油松林地内各层次的氨基酸含量高于连香树林地内相应层次的含量。两林地各层次无机氮含量均超过了氨基酸的含量，并且室内培养30天后无机N的含量仍然高于氨基酸的含量，所以可以认为该区立地条件下无论是在有机分解层还是矿质土层植物吸收利用的氮素仍是以无机N为主。 4 松林下松针凋落物易于累积，这与松针凋落物分解缓慢有关，从而导致松林内养分周转缓慢。通过用不同性质凋落物和灌丛地土壤构建微生态系统，比较油松、辐射松、连香树、灌丛虎榛子凋落物分解对C、N循环过程的影响，结果显示油松和辐射松针叶凋落物比连香树、虎榛子凋落物分解更慢，减缓了养分循环过程。然而将针叶凋落物与阔叶凋落物混合后，油松和辐射松针叶凋落物的分解加快，C、N元素的循环过程也加速。此结果表明在松林内维持具有高质量凋落物的灌丛植被或在松林内栽植一些阔叶树种如连香树对维持和增进松树人工林的土壤肥力有重大的作用。室内培养的结果还显示添加凋落物后土壤DOC和DON的含量显著增加，表明凋落物是土壤DOM的直接来源。然而不同物种凋落物处理下土壤DOC和DON的含量有所不同，并随时间发生改变。混合凋落物处理下土壤DOC和DON的含量均高于松针凋落物单独处理下土壤DOC和DON的含量。DON是一个主要的水溶性N库，随时间的变化趋势与无机N的变化趋势不同，在土壤N循环过程中起到了中间N库的作用。 As a labile fraction of soil organic matter, dissolved organic matter (DOM) plays a very important role in material cycling of terrestrial ecosystem. The turnover of DOM is now being considered as main components in nutrient cycling. DOM mainly includes dissolved organic carbon (DOC), -nitrogen (DON), -phosphorous (DOP) and –sulfur (DOS). Among these constituents, DOC and DON directly participate in C and N cycling. It is essential to study DOC and DON dynamics and their controlling factors in the areas where no related study has ever been carried out. Study about them can provide data supports on understanding the mechanism of the global C and N cycling. DOC and DON dynamics and their controlling factors have been focused on in the research of C and N cycling of forest ecosystems. Based on forest plantations of Maoxian, Minjiang River in warm temperate zone, soil DOC and DON pool size, their seasonal dynamics, and the correlation between DOC, DON and other nutrients were studied in order to understand the importance of DOC and DON in the study area. Soil DOC and DON dynamics induced by leaf litter decomposition were also studied. The study contributed to comprehensively understanding C and N cycling processes and providing baseline data for including DOC and DON into the indices system of evaluating nutrient conditions. The results were as follows: 1 Several different plant communities under brown soil and Cinnamon soil were chosen as sampling plots. The contents and features of soil DOC and DON were evaluated. The results showed that DOC and DON were present under the two soil types. DOC contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 111.96 mg kg-1to 159.95 mg kg-1, and 69.02 mg kg-1 to 100.84 mg kg-1. DON contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 11.88 mg kg-1to 23.08 mg kg-1, and 4.70 mg kg-1 to 10.77 mg kg-1. Free amino acid contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 0.84 mg kg-1to 1.66 mg kg-1, and 0.39 mg kg-1 to 0.73 mg kg-1. Significant correlations were found between DOC, DON and some nutrient indices, which together reflected soil condition and quality. It was hence essential to study DOC and DON in the study area. 2 Seasonal dynamics of DOC, DON, inorganic N, microbial biomass C and N were studied under Pinus tabulaeformis and Cercidiphyllum japonicum plantation. The results indicated that seasonal dynamics of soil DOC and DON under the two plantations performed similar change pattern, with the highest values in autumn. The seasonal dynamics of soil DOC and DON were mainly influenced by the litterfall. However, biotic factors such as soil microbial activities and abiotic factors such as precipitation and temperature also controlled the dynamics of soil DOC and DON. The seasonal dynamic of DON was different from that of NO3--N and NH4+-N, which showed that the behavioral differences between DON and inorganic nitrogen. And hence, it was proposed to include DON into soil N cycling in the study area. 3 Amino acid dynamics in Oa and topsoil (0-10 cm) under P. tabulaeformis and C. japonicum plantation were studied. The results showed that amino acid content in Oa was significantly higher than that in mineral soil. At each sampling time, significantly higher amino acid contents were found in P. tabulaeformis plantation than in C. japonicum plantation. The content of inorganic nitrogen was much higher than the content of amino acid in each sampling layer at each sampling time. After a 30-days laboratory incubation the content of amino acid was still lower than the content of inorganic nitrogen. The results implicated that the form of N absorbed by plants in these study sites were mainly inorganic nitrogen. 4 Usually needle litter is more resistant to decomposition, which leads to needle litter accumulation in pure coniferous stands and slows down the rate of nutrient circulation. By constructing microcosms with local shrubland soil and containing the four single-species (P. tabulaeformis, P. radiata, C. japonicum, Ostryopsis davidiana) litters, the decomposition rates and related C and N dynamics of needle litters and broadleaved litters during the early stage were compared. The results showed that the decomposition rates of pine needles were lower than those of broadleaved litters, which descended C and N cycling processes. However, the presence of C. japonicum or O. davidiana litter into pine needles increased the decomposition rates of pine needles and also dramatically promoted C and N cycling processes. It should be appropriate for plantation managers to consider C. japonicum as an ameliorative species or remain O. davidiana in pine plantations to improve soil conditions and help maintain soil fertility. The laboratory incubation still showed that DOC and DON contents in all litter-amended treatments were significantly higher than no litter-amended treatment, which proved that litter could be a direct source of DOM in soils. Different species litters induced different soil DOC and DON contents, which correspondingly changed over time. DOC and DON contents in mixed litter treatments were higher than those in pine needle litter treatments. As a major soluble N pool, DON developed a different changing pattern over time compared with inorganic N and played a role of interim N pool in soil N cycling.

瑞香狼毒（Stellera chamaejasme L.）对高寒草甸系统生长和非生长季节碳氮循环的影响及其花粉化感效应

瑞香狼毒（Stellera chamaejasme L.）是瑞香科（Thymelaeaceae）狼毒属的一种多年生野草，有毒。据调查，从20 世纪60 年代开始至今，狼毒在青藏高原东缘的高寒草甸上不断蔓延、密度不断变大，在一些地段甚至成为优势物种。有关狼毒在高寒草甸蔓延的生态系统效应的研究尚未见报道。本文从系统碳、氮循环的角度，分别研究狼毒在生长和非生长季节对高寒草甸生态系统的影响。同时，从花粉化感的角度，深入研究狼毒对当地同花期物种有性繁殖的影响。系统地研究高寒草甸生态系统物质循环过程，特别是非生长季节微生物和土壤碳氮库的动态变化，有助于揭示狼毒在系统物质循环方面的“物种效应”以及这种效应的季节变化，为丰富有关高海拔生态系统，特别是其非生长季的物质循环的科学理论做出贡献。同时，碳氮循环和花粉化感的研究还有助于深刻地理解狼毒作为一种入侵性很强的杂草的特殊的蔓延机制，从而为狼毒的有效防治、高寒草甸的科学管理提供依据。 针对狼毒在青藏高原高寒草甸上蔓延的生态系统碳氮循环方面的影响，开展以下2 方面的研究：（1）在生长季，研究松潘县尕米寺附近（北纬32°53'，东经103°40'，海拔3190 m）的两种地形（平地和阳坡）条件下狼毒对土壤碳氮循环影响及可能的原因。狼毒和其它几个主要物种（圆穗蓼（Polygonummacrophyllum D. Don var. Macrophyllum），草地早熟禾（Poa pretensis L.），四川嵩草（Kobresia setchwanensis Hand.-Maizz.），鹅绒委陵菜（Potentilla anserina L.var. anserine）和鳞叶龙胆（Gentiana squarrosa Ledeb.）的地上凋落物产量以及地上凋落物和根的化学组成被测量。在有-无狼毒斑块下，各种土壤的库（比如，铵态氮、硝态氮、无机磷和微生物生物量）和周转率（包括净矿化、净硝化、总硝化、反硝化和微生物呼吸速率）被测量和比较。（2）在非生长季节，尤其是春季冻融交替期，选取了两个研究地点——尕米寺和卡卡沟（北纬32°59'，东经103°41'，海拔3400 m），分别测定有狼毒和无狼毒斑块下土壤微生物生物量碳和氮、可溶性有机碳和氮以及铵态氮和硝态氮的动态变化。同时，分别在上述两个地点有-无狼毒的样地上，研究6 个主要物种（狼毒、圆穗蓼、草地早熟禾、四川嵩草、鹅绒委陵菜和鳞叶龙胆）从秋季开始、为期1 年的凋落物分解过程。 针对狼毒花粉化感对同花期其它物种可能的花粉化感作用开展以下工作：在实验室中，用一系列浓度的狼毒花粉水浸提液对与它同花期的其它物种以及自身花粉进行测试，测定花粉萌发率；在野外自然条件下的其它物种的柱头上施用上述浓度的狼毒花粉水浸提液，观测种子结实率，同时，观察狼毒花粉的种间花粉散布数量。 生长季节的研究结果表明，狼毒地上凋落物氮含量比其它几个主要物种更高，而木质素-总氮比更低。狼毒显著地增加其斑块下表层土壤中有机质的含量，而有-无狼毒的亚表层土壤在有机碳和总磷方面没有显著差异。狼毒表土中硝态氮含量在平地和阳坡比无狼毒土壤分别高113%和90%。狼毒表土中微生物生物量碳和氮量显著高于无狼毒表土。无论是平地还是阳坡，狼毒土壤的总硝化和微生物呼吸速率显著高于无狼毒土壤；而它们的反硝化速率只在平地有显著的差异。狼毒与其它物种间地上凋落物的产量和质量的差异可能是导致有-无狼毒土壤碳氮循环差异的原因。我们假设，狼毒可能通过增加贫氮生态系统土壤中的有效氮含量提高其入侵能力。 非生长季的研究结果表明，在青藏高原东缘的高寒草甸上，土壤微生物生3物量在11 月的秋-冬过渡期达到第一个峰值；在春季的冻融交替期，微生物生物量达到第二个峰值后又迅速降低。无机氮以及可溶性有机碳氮与微生物生物量有相似的变化过程。微生物碳氮比呈现显著的季节性变化。隆冬季节的微生物生物量碳氮比显著高于生长旺季的微生物碳氮比。这种变化可能暗示冬、夏季微生物的群落组成和对资源的利用有所不同。有-无狼毒斑块下土壤微生物和土壤碳、氮库一般只在秋-冬过渡期有显著差异，有狼毒土壤微生物生物量和土壤碳、氮库显著高于无狼毒土壤；而在之后的冬季和春季没有显著差异。所有6 个物种凋落物在非生长季分解率为24%-50%，均高于生长季的10%到30%。其中在秋-冬过渡期，凋落物开始埋藏的两周时间内，分解最快，达10%-20%。不同物种凋落物全年的分解率和分解过程有显著差异。圆穗蓼在全年的分解都较缓慢（非生长季26%，生长季15%），草地早熟禾和四川嵩草等全年的分解速率比较均匀（非生长季和生长季均为30%，非生长季略高），而狼毒在非生长季分解较快（约50%），而在接下来的生长季分解变得缓慢（约12%）。所有物种的凋落物氮含量在非生长季下降，而在随后的生长季上升。 实验室的花粉萌发试验证明，狼毒花粉对自身花粉萌发没有自毒作用，而其它受试的所有物种（圆穗蓼，秦艽（Gentiana macrophylla Pall. var. fetissowii），湿生扁蕾（Gentianopsis paludosa (Hook. f.) Ma var. paludosa），鳞叶龙胆，椭圆叶花锚（Halenia elliptica D. Don var. elliptica），蓝钟花（Cyananthus hookeri C. B.Cl. var. grandiflorus Marq.），小米草（Euphrasia pectinata Ten.），川西翠雀花（Delphinium tongolense Franch.），高原毛茛（Ranunculus tanguticus (Maxim.)Ovcz. var. tanguticus）和鹅绒委陵菜）的花粉萌发率随着狼毒花粉浸提液浓度的增加呈显著的非线性降低。大约3 个狼毒花粉的浸提液就可以抑制受试的多数物种的50%的花粉萌发。在鳞叶龙胆和小米草柱头上狼毒花粉的数量分别为5.76 个和3.35 个。狼毒花粉散布数量的差异最可能的原因在于是否有共同的传粉昆虫。花的形状（辐射对称VS 左右对称）、植株或花的密度以及花期重叠性可以部分解释这种差异。在野外试验中，我们发现6 个物种（秦艽、湿生扁蕾、鳞叶龙胆、椭圆叶花锚、蓝钟花和小米草）的种子结实率随狼毒花粉浸提液浓度的增加呈显著的非线性降低。鳞叶龙胆和小米草柱头上狼毒花粉的数量（分别是5.76 个和3.35 个）分别达到了抑制它们63%和55%种子结实率的水平。因此，狼毒对鳞叶龙胆和小米草可能存在明显的花粉化感抑制作用。狼毒周围的物种可能通过花期在季节或昼夜上的分异避免受到狼毒花粉化感的影响或者通过无性繁殖来维持种群繁衍，因此狼毒通过花粉化感作用对其周围物种繁殖的影响程度还需要进一步地研究。如果狼毒的花粉化感抑制作用确实存在，那么它可能成为一种自然选择压力，进而影响物种的进化。 Stellera chamaejasme L., a perennial toxic weed, has emerged and quicklydominated and spread in the high-frigid meadow on the eastern Tibetan Plateau ofChina since the 1960s. In the present study, effects of S. chamaejasme on carbon andnitrogen cycles on the high-frigid meadow on the eastern Qinghai-Tibetan Plateau ingrowing and non-growing season, and its pollen allelopathic effects on the sympatricspecies were determined. The present study that focused on carbon and nitrogencycles, especially on microbial biomass and pools of carbon and nitrogen innon-growing season, could profoundly illuminate plant-species effects on carbon andnutrient cycles and its seasonal pattern and help to understand spread mechanism ofS. chamaejasme as an aggressive weed. The present study also contributed to furtherunderstand carbon and nutrient cycles on alpine regions in non-growing season andprovide a basis on weed control of S. chamaejasme and scientific management in thehigh-frigid ecosystem. Effects of S. chamaejasme on carbon and nitrogen cycles on the high-frigidmeadow on the eastern Qinghai-Tibetan Plateau were determined. The study couldbe divided into two parts. (1) In the growing season, we quantified the effects of S.chamaejasme on carbon and nitrogen cycles in two types of topographic habitats, theflat valley and the south-facing slope, where S. chamaejasme was favored to spreadlitter and root were measured to explain the likely effects of S. chamaejasme on soilcarbon and nutrient cycles. The sizes of various soil pools, e.g. nitrite, ammonium,inorganic phosphorus and microbial biomass, and turnover rates including netmineralization, gross nitrification, denitrification and microbial respiration weredetermined. (2) In the non-growing season study, microbial biomass carbon andnitrogen, soluble organic carbon and nitrogen, ammonium and nitrate weredetermined through the non-growing season, especially in the processes offreeze-thaw of spring in two high-frigid sites, i.e. Kaka valley and Gami temple, onthe eastern Qinghai-Tibetan Plateau. Meanwhile, litter decomposition of six commonspecies, including Stellera chamaejasme L., Polygonum macrophyllum D. Don var.Macrophyllum, Poa pretensis L., Kobresia setchwanensis L., Potentilla anserina L.var. anserine and Gentiana squarrosa Ledeb., were also examined under theabove-mentioned experimental design through one whole-year, which began in theautumn in 2006. In the study of pollen allelopathy, several work, including in vitro study oneffects of extract of pollen from S. chamaejasme on sympatric species and pollenfrom itself, field experiments on effects of pollen extract with the same regime ofconcentrations on seed set and field observation on heterospecific pollen transfer ofS. chamaejasme to six of those sympatric species has been done. The results in the growing season showed that aboveground litter of S.chamaejasme had higher tissue nitrogen and lower lignin: nitrogen ratio than thoseco-occurring species. S. chamaejasme significantly increased topsoil organic matter,whereas no significant differences were found for organic C and total P in subsoilbetween under-Stellera and away-Stellera locations. The nitrate in Stellera topsoilwas 113% and 90% higher on the flat valley and on the south-facing slope,respectively. Both microbial biomass C and N were significantly higher in Stelleratopsoil. Gross nitrification and microbial respiration were significantly higher inStellera topsoil both on the flat valley and on the south-facing slope, whereassignificant differences of denitrification were found only on the flat valley. Thedifferences in the quantity and quality of aboveground litter are a likely mechanismresponsible for the changes of soil variables. We assumed that S. chamaejasme couldenhance their spread by increasing nutrient availability in N-deficient ecosystems. The results in the non-growing season showed that microbial biomass achievedthe first summit in late autumn and early winter on the eastern Qinghai-TibetanPlateau. In the stages of freeze-thaw of spring, microbial biomass firstly achieved thesecond summit and subsequently sharply decreased. Inorganic nitrogen, solubleorganic carbon and nitrogen had a similar dynamics with that of microbial biomass.Ratio of microbial biomass carbon and nitrogen had an obviously seasonal pattern.The highest microbial C: N were in the non-growing season, which weresignificantly higher than those in the growing season. The seasonal pattern inmicrobial biomass C: N suggested that large changes in composition of microbialpopulation and in resources those used by microbes between summer and winter.Generally, microbial biomass and pools size of carbon and nitrogen in Stellera soilwere significantly higher than those under adjacent locations in late autumn andearly winter, but there were not significant differences in winter and in spring. Litterof all the focal species (Stellera chamaejasme L., Polygonum macrophyllum D. Donvar. Macrophyllum, Poa pretensis L., Kobresia setchwanensis Hand.-Maizz.,Potentilla anserina L. var. anserine and G. squarrosa Ledeb.) decomposed about24%-50% in the non-growing season, which were higher than those in the growingseason (ranged from 10% to 30%). Litter decomposed 10%-20% within the first twoweeks in late autumn and early winter. Significant differences in the whole-yeardecomposition rate and in the processes of decomposition were found among species.Polygonum macrophyllum decomposed slowly through the whole year (26% and15% in the non-growing season and in the growing season, respectively). Certainspecies, such as P. pretensis L. and K. setchwanensis, decomposed at a similar rate(30% both in the non-growing and in the growing season, slightly higher in the8growing season than those in the growing season), whereas S. chamaejasmedecomposed more rapidly (about 50%) in the non-growing season and subsequentlydecomposition became slow (about 12%) in the growing season. Litter nitrogencontents of all the focal species firstly decreased in the non-growing season and thenincreased in the growing season. In vitro experiments of pollen allelopathy, the results showed that pollen from S.chamaejasme was not autotoxic, whereas pollen germination in all the sympatricspecies (Polygonum macrophyllum D. Don var. Macrophyllum, Gentianamacrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Ma var. paludosa,Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica, Cyananthushookeri C. B. Cl. var. grandiflorus Marq., Euphrasia pectinata Ten., Delphiniumtongolense Franch., Ranunculus tanguticus (Maxim.) Ovcz. var. tanguticus andPotentilla anserina L. var. anserina) decreased nonlinearly as the increasingconcentrations of extract of pollen from S. chamaejasme. Pollen Extract of threepollens from S. chamaejasme generally inhibited 50% pollen germination of most ofthe focal species. 5.76 and 3.35 pollens from S. chamaejasme were observed in fieldon stigmas of G. squarrosa and E. pectinata, respectively. Differences inheterospecific pollen transfer of S. chamaejasme could be attributed to the primaryreason whether they shared common pollinators. Flower morphology (e.g.zygomorphic or actinomorphic), plant or floral density and concurrence in floweringphonologies could explain, in part, the differences in heterospecific pollen transfer.In field experiments, the results showed that seed set in six sympatric species(Gentiana macrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Mavar. paludosa, Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica,Cyananthus hookeri C. B. Cl. var. grandiflorus Marq. and Euphrasia pectinata Ten.)decreased nonlinearly as the increasing concentrations of extract of pollen from S.chamaejasme. According to the nonlinear curves, the amounts of pollens from S.chamaejasme on stigmas of G. squarrosa and of E. pectinata (i.e. 5.76 grains and3.35 grains, respectively) could reduce 63% and 55% seed set of G. squarrosa and ofE. pectinata, respectively. Thus, allelopathic effects of S. chamaejasme on G.squarrosa and E. pectinata could be realistic. The sympatric species of S.chamaejasme could avoid pollen allelopathy of S. chamaejasme to sustainthemselves. This highlights the need to study how much pollen allelopathy in S.chamaejasme influences the sympatric species through divergence in seasonal ordiurnal flowering phonologies or through asexual reproduction. If pollen allelopathyin S. chamaejasme was confirmed, it could be as a pressure of natural selection andthus play an important role in species evolution.

干旱胁迫与施N条件下白刺花（Sophora davidii）幼苗生长及其适应机制

干旱环境常常由于多变的降水事件和贫瘠土壤的综合作用，表现出较低的生产力和较低的植被覆盖度。全球性的气候变暖和人类干扰必将使得干旱地区缺水现状越来越严竣。贫瘠土壤环境中已经很低的有效养分含量也将会随着干旱的扩大而越来越低。干旱与半干旱系统中不断加剧的水分与养分的缺失将严重限制植物的生长和植被的更新，必然会使得已经恶化的环境恶化速率的加快、恶化范围的加大。如何抑制这种趋势，逐步改善已经恶化的环境是现在和将来干旱系统管理者面临的主要关键问题。了解干旱系统本土植物对未来气候变化的适应机制，不仅是植物生态学研究的重要内容，也对人为调节干旱环境，改善干旱系统植被条件，提高植被覆盖度具有重要的实践意义。 本研究以干旱河谷优势灌木白刺花（Sophora davidii）为研究对象，通过两年大棚水分和施N控制实验和一个生长季野外施N半控制实验，从植物生长－生理－资源利用以及植物生长土壤环境特征入手，系统的研究了白刺花幼苗生长特性对干旱胁迫和施N的响应与适应机制，并试图探讨施N是否可调节干旱系统土壤环境，人工促进干旱条件下幼苗定居，最终贡献于促进植被更新实践。初步研究结论如下： 1）白刺花幼苗生长、生物量积累与分配以及水分利用效率对干旱胁迫和施N处理的适应白刺花幼苗株高、基径、叶片数目、叶面积、根长、生物量生产、相对含水量和水分利用效率随着干旱胁迫程度的增加而明显降低，但地下部分生物量比例和R/S随着干旱胁迫程度的增加而增加。轻度施N处理下幼苗株高、基径、叶片数目、叶片面积和生物量生产有所增加。但重度施N处理下这些生长指标表现出微弱甚至降低的趋势。严重干旱胁迫条件下，幼苗叶面积率、R/S、相对含水量和水分利用效率也以轻度施N处理为最高。 2）白刺花幼苗叶片光合生理特征对干旱胁迫和施N处理的适应叶片光合色素含量和叶片光合效率随着干旱胁迫程度的增加而显著降低，并且PS2系统在干旱胁迫条件下表现出一定程度的光损害。但是比叶面积随着干旱胁迫程度的增加而增加。在相对较好水分条件下幼苗净光合速率的降低可能是因为气孔限制作用，而严重干旱胁迫条件下非气孔限制可能是导致幼苗叶片光合速率下降的主要原因。叶片叶绿素含量、潜在光合能力、羧化效率、光合效率以及RUBP再生能力等在施N处理下得到提高，并因而改善干旱胁迫条件下光合能力和效率。虽然各荧光参数对施N处理并无显著的反应，但是干旱胁迫条件下qN和Fv/Fm在轻度施N处理下维持相对较高的水平，而两年连续处理后在严重干旱胁迫条件下幼苗叶片光合效率受到重度施N处理的抑制，并且Fv/Fm和qN也在重度施N处理下降低。 3）白刺花幼苗C、N和P积累以及N、P利用效率对干旱胁迫和施N处理的适应白刺花幼苗C、N和P的积累，P利用效率以及N和P吸收效率随干旱胁迫程度的增加而显著降低，C、N和P的分配格局也随之改变。在相同水分处理下，C、N和P的积累量、P利用效率以及N和P吸收效率在轻度施N处理下表现为较高的水平。然而，C、N和P的积累量和P利用效率在重度施N处理下不仅没有表现出显著的正效应，而且有降低的趋势。另外，在相同水分条件下白刺花幼苗N利用效率随着施N强度的增加而降低。 4）白刺花幼苗生长土壤化学与微生物特性对干旱胁迫和施N的适应白刺花幼苗生长土壤有机C、有效N和P含量也随干旱胁迫程度的增加而明显降低。干旱胁迫条件下土壤C/N、C/P、转化酶、脲酶和碱性磷酸酶活性的降低可能表明较低的N和P矿化速率。尽管微生物生物量C、N和P对一个生长季干旱胁迫处理无显著反应，但微生物生物量C和N在两年连续干旱胁迫后显著降低。土壤有机C和有效P含量在轻度施N处理下大于重度施N处理，但是有效N含量随着施N强度的增加而增加。微生物生物量C和N、碱性磷酸酶和转化酶活性也在轻度施N处理下有所增加。但是碱性磷酸酶活性在重度施N处理下降低。 5）野外条件下白刺花幼苗生长特征及生长土壤生化特性对施N的适应植物生长、生物生产量、C的固定、N、P等资源的吸收和积累、其它受限资源的利用效率（如P）在轻度施N处理下均有所增加，但N利用效率有所降低。幼苗生物生产量及C、N和P等资源的分配格局在轻度施N处理下也没有明显的改变。白刺花幼苗叶片数目、生物生产量和C、N、P的积累量在重度施N处理下虽然也相对于对照有所增加，但幼苗根系长度显著降低。生物量及资源（生物量、C、N、P）在重度施N处理下较多地分配给地上部分（主要是叶片）。另外，土壤有机C、全N和有效N含量随外源施N的增加而显著增加，土壤pH随之降低，但土壤全P含量并无显著反应。其中有机C含量和有效P含量以轻度施N处理最高。微生物生物量C、N和P在轻度施N处理下也显著增加，而微生物生物量C在重度施N处理下显著降低。同时，转化酶、脲酶、碱性磷酸酶和中性磷酸酶活性在施N处理下也明显的提高，但酸性磷酸酶和过氧化氢酶活性显著降低，其中碱性磷酸酶和中性磷酸酶活性以轻度施N处理最高。 综合分析表明，干旱河谷水分和N严重限制了白刺花幼苗的生长。施N不能完全改变干旱胁迫对白刺花幼苗的抑制的作用，但是由于施N增加土壤N有效性，改善土壤一系列生物与化学过程，幼苗的生长特性也对施N表现出强烈的反应，表现为植物结构与资源分配格局的改善，植物叶片光合能力与效率的提高，植物生长以及利用其他受限资源（如水分和P）的效率的增加，致使植物自身生长及其生长环境在干旱环境下得到改善。但是过度施N不仅不能起到改善干旱胁迫下植物生长环境、促进植物生长的作用，反而在土壤过程以及植物生长过程中加重干旱胁迫对植物的伤害。因此，建议在采用白刺花作为先锋种改善干旱河谷系统环境的实践中，可适当施加N以改善土壤环境，调节植物利用与分配资源的效率，促进植物定居，得到人工促进种群更新的目的。但在实践过程中也要避免过度施N。 Arid regions of the world are generally noted for their low primary productivity which is due to a combination of low, unpredictable water supply and low soil nutrient concentrations. The most serious effects of global climate change and human disturbances may well be those which related to increasing drought since drought stress has already been the principal constraint in plant growth. The decline in total rainfall and/or soil water availability expected for the next decades may turn out to be even more drastic under future warmer conditions. Nevertheless, water deficit is not the only limiting factor in arid and semiarid environments. Soils often suffer from nutrient (especially N and P) deficiencies in these ecosystems, which can also be worsened by climate change. How to improve the poor soil quality and enhance the vegetation coverage is always the problem facing ecosystem managers. The adaptive mechanisms of native plant to future climate change is always the focus in plant ecology, it also plays important roles in improving vegetation coverage by manual controlled programmes. Sophora davidii is a native perennial shrub of arid valleys, which is often predominant on eroded slopes and plays a vital role in retaining ecological stability in this region. It has been found that S. davidii was better adapted to dry environment than other shrubs, prompting its use for re-vegetation of arid lands. A two-years greenhouse experiment and a field experiment were conducted in order to understand the adaptation responses of Sophora davidii seedlings to different water and N conditions, and further explore if additional N supply as a modified role could enhance the adaptation ability of S. davidii seedlings to dry and infertile environment. Two-month old seedlings were subjected to a completely randome design with three water (80%, 40% and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg-1 soil) regimes. Field experiment was arranged only by three N supplies in the dry valley. 1) The growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings in respond to drought stress and N supply Seedlings height, basal diameter, leaf number, leaf area, root length, biomass production, relative water content (RWC) and WUE were decreased with increase of drought stress. An increase in below-ground biomass was observed indicating a higher root/shoot ratio (R/S) under drought stress conditions. Low N supply increased seedlings height, basal diameter, leaf number, leaf area, and biomass production, but decreased root length. In contrast, these growth characteristics showed little or negative effect to high N supply treatment. Leaf percentages increased with increase of N supply, but fine root percentages decreased. In addition, Low N supply rather than the other two N treatments increased leaf area ratio (LAR), leaf/fine root mass ratio (L/FR), R/S and RWC under severe drought stress (20%FC), even though these parameters could increase with the high N supply treatment under well-watered condition (80%FC). Moreover, Low N supply also increased WUE under three water conditions, but high N supply had little effect on WUE under drought stress conditions (40%FC and 20%FC). 2) Leaf gas exchange and fluorescence parameters of Sophora davidii seedlings in respond to drought stress and N supply Leaf area (LA), photosynthetic pigment contents, and photosynthetic efficiency were decreased with increase of drought stress, but specific leaf area (SLA) increased. Photodamage in photosystem 2 (PS2) was also observed under drought stress condition. The decreased net photosynthetic rate (PN) under relative well-watered water conditions might result from stomatal limitations, but the decreased PN under other hand, photosynthetic capacity by increasing LA, photosynthetic chlorophyll contents, Pnmax, CE, Jmax were increased with increase N supply, and photosynthetic efficiency was improved with N supply treatment under water deficit. Although N supply did a little in alleviating photodamages to PS2 caused by drought stress, low N supply enhanced qN and kept relative high Fv/Fm under drought stress condition. However, high N supply inhibited leaf photosynthetic efficiency, and declined Fv/Fm and qN under severe drought stress condition after two year continues drought stress and N supply. 3) Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in respond to drought stress and N supply C, N and P accumulation, NUE , N and P uptake efficiency (NUtE and NUtE ) P N P were decreased with increase of drought stress regardless of N supply. On the other hand, the S. davidii seedlings exhibited strong responses to N supply, but the responses were inconsistent with the various N supply levels. Low N supply rather than the other two N treatments increased C, N and P accumulation, improved NUEP, NUtE and NUtE under corresponding water condition. In contrast, high N supply N P did few even depressed effects on C, N and P accumulation, and NUEP, although NUtEN and NUtEP could increase with high N supply under corresponding water conditions. Even so, a decrease of NUEN was observed with increase of N supply under corresponding water conditions. 4) Soil microbial and chemical characters in respond to drought stress and N supply The content of soil organic C, available N and P were decreased with increase of drought stress. Decreases in C/N and C/P, and invertase, urea and alkaline phosphatase activity were also observed under drought stress conditions, indicating a lower N and P mineralization rate. Although microbial biomass C, N and P showed slight responses to drought stress after one growth period treatment, microbial biomass C and N were also decreased with increase of drought stress after two year continuous treatment. The content of soil organic C and available P showed the stronger positive responses to low N supply than which to high N supply, although than the other two N treatments increased microbial biomass N and invertase activity under severe drought stress condition, even though invertase activity could increase with high N supply treatment under relative well-water conditions. Moreover, low N supply treatment also increased C/P and alkaline phosphatase activity which might result from higher P mineralization, but high N supply did negative effects on alkaline phosphatase activity. 5) The growth characteristics of Sophora davidii seedlings and soil microbial and chemical characters in respond to N supply under field condition Low N supply facilitated seedlings growth by increasing leaf number, basal diameter, root length, biomass production, C, N and P accumulation and absorption, and enhancing the use efficiency of other limited resources as P. Compared to control, however, low N supply did little effect on altering biomass, C, N and P portioning in seedlings components. On the contrary, high N supply treatment also increased leaf number, biomass and C, N and P accumulation relative to control, but significantly decreased root length, and altered more biomass and resources to above-ground, which strongly reduced the ability of absorbing water under drought condition, and thus which might deep the drought stress. In addition, N supply increased soil C, N and available N content, but declined pH and showed little effects on P content. Low N supply showed higher values of soil C and available P content. Low N supply also increased microbial biomass C, N and P, although high N supply decreased microbial biomass C. N supply significantly enhanced soil invertase, urea, alkaline and neutral phosphratase activity, while declined acid phosphratase and catalase activity. Low N supply exhibited higher alkaline and neutral phosphratase activity compared to the others. The results from this study indicated that both drought and N limited the growth of S. davidii seedlings and their biomass production. Regardless of N supply levels, drought stress dramatically reduced the seedlings growth and biomass production. Although plant growth parameters, including basal diameter, height, leaf number, and biomass and their components were observed to be positive responses to low N supply, N supply alone can not alter the diminishing tendency which is caused by drought. available N content increased with increase N supply. In addition, low N supply rather These findings imply that drought played a primary limitation role and N was only the secondary. Even so, appropriate N supply was seemed to enhance the ability that S. davidii seedlings adapted to the xeric and infertile environment by improving soil processes, stimulating plant growth, increasing recourses accumulation, enhancing use efficiency of other limited resources, and balancing biomass and resources partitioning. Appropriate N supply, therefore, would be recommended to improve S. davidii seedling establishment in this region, but excess N supply should be avoided.

Short-term effect of targeted placements of sheep excrement on grassland in inner mongolia on soil and plant parameters

Grazing animal excrement plays an important role in nutrient cycling and redistribution in grazing ecosystems, due to grazing in large areas and return in small areas. To elucidate the changes to the soil and pasture caused by sheep urine, fresh dung, and compost patches, a short- term field experiment using artificially placed pats was set up in the autumn of 2003 in the Inner Mongolian steppe. Urine application significantly increased soil pH during the first 32 days in soil layers at depths of both 0 - 5 cm and 5 - 15 cm. Rapid hydrolysis of urea gave large amounts of urine- nitrogen ( N) as ammonium ( NH4+) in soil extracts and was followed by apparent nitrification from day 2. Higher inorganic N content in the urine- treated soil was found throughout the experiment compared with the control. No significant effects of sheep excrement on soil microbial carbon ( C) and soil microbial N was found, but microbial activities significantly increased compared with the control after application of sheep excrement. Forty- six percent of dung- N and 27% of compost- N were transferred into vegetation after the experiment. The results from this study suggest that large amounts of nutrients have been lost from the returned excrement patches in the degraded grassland of Inner Mongolia, especially from sheep urine- N.