中国植物功能型-生物群区的划分

气候在大尺度上决定着植被的分布、结构和组成，植被结构和生理状态的改变可以通过改变植被的反射率、粗糙度以及水分通量进而影响气候，这样形成了气候一植被的相互作用。在植被一气候相互作用的研究中，植物功能型是重要的概念和方法，它可以在详尽描述植被生物物理和生理特征的同时，有效削减植被的复杂性。植物功能型的概念和方法已经在植物群落、生态系统的复杂性和功能、古植被和古气候研究，以及陆面过程模型和动态全球植被模型中得到了广泛的应用。但是针对我国植被-气候的相互作用和区域尺度的全球变化研究，还需要一套特定的植物功能型．生物群区体系。 　　本论文根据我国植被生态学和植被分类的研究背景，结合植被．气候相互作用和区域全球变化研究的需要，提出了一套适宜于中国的植物功能型．生物群区划分方案。首先，根据中国植被和气候特征，筛选并确定了影响植被生物物理和生理属性以及植被分布的6个关键的植物功能特征：然后，根据这6个特征，对植物进行功能型划分，得到了29类植物功能型：再根据我国植被的实际情况和研究需要，选定了其中的18类作为我国的植物功能型。这套功能型包括了7类‘树’功能型，6类‘灌木’功能型和5类‘草’功能型，其中含有4类高寒植物功能型，专门用于描述青藏高原的植被分布，并根据需要设置了2类‘裸地，功能型。 　　根据我国气候一植被分布定量关系的相关研究以及BIOME1和Box体系的研究结果，选定7个环境变量作为限制我国植物功能型分布的关键气候因子：最冷月平均气温、最暖月平均气温、大于50C的有效生长积温、大于OºC的有效生长积温、Priestley-Taylor系数（实际蒸散与潜在蒸散的比值）、降水量、最暖月和最冷月平均气温之差。采用半峰宽法初步确定每个植物功能型的环境限定因子取值范围。并根据这套植物功能型及其环境参数建立了适宜于我国的生物群区体系，从而得到了我国的植物功能型-生物群区体系(the Chinese Plant functional Types and Biomes，CNPB)。 为了验证这套植物功能型-生物群区体系，将BIOME1和中国的植物功能型生物群区体系(CNPB)对中国植被在当前气候条件和未来气候情景下分布的模拟结果进行了比较。结果表明，这套体系可以更有效地模拟中国植被在当前和未来气候条件下的分布，特别是对青藏高原植被描述的详细程度有实质性的提高。

京北农牧交错区植物功能型研究

植物功能型的研究是全球变化研究当中最重要的创新之一，作为新的研究手段，已经被广泛用来研究全球变化的影响。而关于不同植物功能型对全球变化的生理生态反应、植物功能型对全球变化的适应机制及植物功能型演变在全球变化中的作用等方面的研究报道却很少，相关方面的研究尚处于起步阶段，而这些恰恰是植物功能型研究的基本点和最终目的，尤其对于全球变化预测模型的研究具有深远的意义。因此深入开展区域环境或全球变化下植物功能型的变化和适应机制研究能为最大程度地获得全球变化对未来影响提供重要信息。 　　本研究中我们对京北农牧交错区的植物功能型（光合功能型和形态功能型）进行了研究。利用稳定性同位素判定法对锡林郭勒草原四种不同草地类型植物功能型进行了分析研究，在调查的125种维管植物中有4种C4植物，其中禾本科和藜科各有2种，C4植物约占该地区调查植物总数的3%，C3 植物为94%、CAM植物为3%。研究中发现利用稳定性同位素技术判定CAM类型存在一定的局限性，需要采用辅助实验进行补充。研究结果还表明自东向西，在草甸草原－典型草原－荒漠草原这一系列草原类型中随着降雨量的逐渐降低，植物种类组成逐渐降低，但C4/C3的比例却有升高的趋势，说明植物光合类型的组成与水分梯度相关。在相同的环境梯度下，多年生高禾草和多年生杂类草呈现显著减少的趋势;而一年生杂类草，多年生矮禾草、灌木和鳞茎类植物呈现显著增加的趋势。这些结果为我们预测该地区植物功能型对全球变化影响的响应提供了依据。 　　在北京北部农牧交错区对不同生境类型的植物功能型进行了研究，该区域有C4植物68种。具有C4 植物的科相对集中，主要分布在禾本科（29属43种）、莎草科（4属16种）和藜科（3属5种）。菊科、豆科和蔷薇科这3大科中没有发现C4 植物。C4植物约占该地区调查植物总数的9%，C3 植物为89%、CAM植物为2%。多年生高禾草和多年生杂类草的比例无论从生长条件良好的草地到沙地，还是从草地到弃耕干扰地都呈现显著下降的趋势，而一年生杂类草和一年生禾草的变化却呈相反的变化趋势。C4植物中一年生植物占的比例最大，而且各个生境中C4植物在一年生植物的比例均超过30%，这说明该地区各个生境植被都已有相当程度的退化。上述结果证明植物功能型能在不同生境尺度上反映土地利用的变化。 　　对典型农牧交错区（多伦县）植物功能型研究表明，在该地区调查的11个群落中有4种C4植物（2科3属），其中禾本科3种、藜科1种。 C4 植物约占该地区调查植物总数的4.5%，C3 植物为95%，CAM植物为0.5%。研究结果表明随着土壤水分含量的降低，C4 功能型以及一年生杂草、一年生禾草、多年生矮禾草对群落水分利用的贡献呈现出显著增加的趋势，而多年生杂类草和多年生高禾草则呈现相反的变化趋势。不同植物功能型的水分利用效率是一个相对稳定的指标，可能成为我们进行更大尺度水分变化分析的有力工具。多年生杂类草和多年生高禾草对维持该地区生态系统生产力以及稳定性具有至关重要的作用，随土壤水分的下降，二者的地位和作用均呈现下降趋势，因此可能导致该地区草地生态系统生产力和稳定性的持续下降。在植物功能型鉴定的基础上，分析了多样性与土壤水分和生产力关系，研究发现植物功能型多样性与土壤水分和生产力相关性更为显著。 　　对3个研究地点植物功能型的比较研究发现，就植物光合功能型而言，随降水量减少和温度降低，单子叶C4光合功能型比例呈现显著下降趋势，而双子叶C4光合功能型却呈现显著增加的趋势;植物形态功能型在区域尺度上没有而呈现出明显的变化趋势。研究表明植物光合功能型对于大尺度下环境因子的变化具有敏感的响应，而植物形态功能型可能更适合在中小尺度下研究环境的异质性以及土地利用方式的变化。 　　

克氏针茅(Stipa krylovii Roshev.)草原植物凋落物分解特性及其对环境变化的响应

凋落物的分解对生物地球化学循环起着重要的作用。本研究以位于中国北方农牧交错区（内蒙古多伦县）的半干旱克氏针茅(Stipa krylovii Roshev.)草原生态系统为背景，针对凋落物分解研究中的几个关键过程和问题进行了探讨。研究内容包括该生态系统中几种常见植物凋落物的分解速率，在分解过程中的养分动态，凋落物的混合效应（即非加性效应）及其机制，全球变化引起的土壤有效N、P和降水增加对凋落物分解的影响及其对草原生态系统碳储量的影响等。 通过对克氏针茅、糙隐子草(Cleistogenes squarrosa Trin.)、双齿葱(Allium bidentatum Fisch.)、野韭(Allium ramosum L. Sp. Pl.)和冰草(Agropyron cristatum Gaertn.)的叶、茎、根凋落物的研究发现，叶和茎的分解速率与凋落物初始N和P含量呈显著正相关，根系的分解速率与初始C/N比呈显著负相关。根系凋落物一般比叶和茎凋落物分解速率快，而且具有更快的养分周转速率。有些凋落物混合在一起分解后对分解速率产生了混合效应（混合凋落物的实际分解速率偏离于基于组分凋落物计算的预期分解速率），同时它们对养分（主要是N、P、Ca、Mg）的固定或释放被延后。混合效应发生与否以及混合效应的方向主要取决于组分凋落物的特点，而与凋落物多样性的高低没有明显的关系。混合凋落物对养分动态的负作用，能降低退化生态系统遭到干扰后养分的损失，而且养分释放时间的延后有助于某些植物更好地生长，有利于退化草原生态系统恢复过程中植被结构的改善。 由于凋落物的质量对分解速率的影响重大，凋落物的化学组成很可能对混合凋落物分解过程中的非加性效应起重要作用，然而迄今为止没有明确的结论。本文研究发现，具有不同初始N、P含量的凋落物（同一物种）混合在一起分解后产生了正的非加性效应，非加性效应的强弱与组分凋落物初始N、P含量的差异大小有关。而且，非加性效应与P含量差异大小的相关性比与N的强，这可能与研究地点土壤中P的含量相对更为缺乏有关。这一研究结果表明，凋落物的化学组成在有些情况下确实与非加性效应有关。 通过施肥和浇水试验，研究了全球变化引起的土壤有效N、P和水分的增加对具有不同生活型和化学组成的两种优势草原植物，即双齿葱和克氏针茅凋落物分解的直接影响。结果发现，向土壤中添加N肥或N、P复合肥，在短期内（100天）加速了这两种植物的分解速率。高质量凋落物（双齿葱）的分解更容易受到土壤水分条件的限制，而低质量凋落物（克氏针茅）的分解对土壤养分有效性更敏感。土壤中有效养分的增加会提高分解凋落物对养分的固定，有利于退化生态系统中养分的保持。双齿葱对内蒙古半干旱典型草原的碳循环和养分动态的贡献，比克氏针茅要更大。研究结果提示人们，对草原生态系统的碳循环和养分动态进行模拟时，需要对不同化学组成的凋落物分别加以考虑。 本文还探讨了土壤有效N长期增加后对两种优势草原植物（克氏针茅和冷蒿(Artemisia frigida Willd)）凋落物分解的综合影响，并进一步区分了由土壤有效N增加引起的凋落物质量改变的间接作用和土壤状况改变的直接作用。结果发现，土壤有效N长期增加后对凋落物的分解速率没有明显影响，但是显著加速了分解凋落物对养分的固定，这可能是由于凋落物质量对分解的促进作用和土壤状况对分解的抑制作用相互抵消的原因所造成的。向土壤中添加N素后，地上部植被生产力不可避免的会提高，尽管短期内会加速凋落物的分解速率，但从长远来看，对凋落物的分解速率不会产生影响，所以估计土壤有效N增加后最终会提高草原生态系统的净碳储量。 通过本研究，对中国北方农牧交错区（内蒙古多伦县）半干旱克氏针茅草原生态系统的养分循环过程有了初步的了解，验证了混合凋落物分解过程中非加性效应发生的可能性，并且证明组分凋落物的初始化学组成对这种非加性效应确实起着重要作用。本研究说明全球变化对不同植物种凋落物分解的影响是不同的，而且因研究时间长短的不同对凋落物分解的影响会发生变化。研究结果为草原生态系统碳和养分循环过程的模拟提供了必要的基础资料，为混合凋落物分解过程中非加性效应发生的可能机制提供了强有力的证据，对在全球变化背景下人们对草原生态系统碳和养分循环过程的认识和模拟有着重要的参考价值。

土地利用方式对内蒙古农牧交错区草地生态系统的影响

陆地生态系统是人类赖以生存与持续发展的基础。由于其巨大的碳储存能力陆地生态系统在全球碳循环过程中发挥着重要作用，尤其是在延缓因大气中CO2浓度升高所引起的气候变化方面作用更为突出。作为世界上最广布的植被类型之一，草地生态系统是陆地生态系统的重要组成部分，其面积占到全球陆地面积的五分之一。中国有着世界第二大的草地生态系统，面积为3.55×108 hm2，约占世界草地面积的6％～8％，其碳储量为世界总碳储量的9％～16％。以内蒙古高原为主体的北方草地，面积约占国土总面积的40％左右，是一种重要的可再生的农牧业资源;作为一种重要的绿色生态屏障，北方草地在调节气候，涵养水源，防风固沙，净化空气和美化环境等方面起着重要作用。由于人口的增长和经济发展的需求，许多土地利用方式呈现在北方农牧交错区。地处内蒙古高原南麓的多伦县属于典型的农牧交错区，良好的气候条件和复杂的人文环境决定了多种土地利用方式在这一地区的并存，为开展不 同土地利用方式对草地生态系统影响的研究提供了天然的研究场所。 本研究选择了多伦县境内的6种土地利用方式作为研究对象，即围封禁牧样地、刈割禁牧样地、自由放牧样地、人工（紫花苜蓿）草地、农田（青贮玉米）和弃耕地，主要研究在这几种不同的土地利用方式影响下，土壤碳氮储量、草地生态系统功能（生产力等）和群落结构的变化。通过对土壤碳氮储量变化的研究发现： (1) 不同土地利用方式因其地上净初级生产力的不同而影响了土壤的碳氮储量，但是高的地上初级生产力并不一定能提高土壤的碳氮储量，如青贮玉米地。这主要是因为土壤碳氮储量与根系和凋落的输入相关。 (2) 不同土地利用方式对植被的浅层（0-20 cm）根系影响较大，继而影响了土壤的碳氮储量。这是因为大约80％的根系碳氮和50％的土壤碳氮储量都集中分布在这一层次。而土壤潜在库容的进一步扩大则要依靠深层根系生物量的增加，以提高深层土壤的碳氮含量，如人工（苜蓿）草地。 (3) 在合理利用方式或者科学管理措施下，草地土壤有机碳储量处于8 kg C m-2到10 kg C m-2之间，比过度退化草地的有机碳储量提高了50％。 (4) 经过短期恢复，土壤的库容量会很快增加，表现出较强的储存功能。因此，通过合理的土地利用方式和管理措施，中国北方农牧交错区可以转变成一个巨大的碳汇。 通过对不同利用方式下草地生态系统的功能、群落结构和物种多样性恢复的研究，得到如下实验结果： (1) 与自由放牧相比，短期（5年）禁牧（围封和刈割）提高了草地的地上净初级生产力和物种数目。围封禁牧和刈割禁牧草地的地上净初级生产力分别比自由放牧草地高出107.3％和50％;而物种数目则分别比自由放牧草地增加了14.8％和25.9％。 (2) 围封和刈割禁牧，尤其是后者，主要增加了非禾本科草本植物在群落中所占的比例;而在自由放牧草地，灌木和半灌木种群在群落中占有绝对优势。 (3) 物种多样性和生产力间的关系因外部干扰程度的不同而变化。在轻度干扰（围封禁牧）和中度干扰（刈割禁牧）下，生产力和物种多样性间呈指数正相关关系;而在重度干扰（自由放牧）下，群落地上净初级生产力随着物种多样性的增加呈指数递减的趋势。 通过对不同土地利用（或干扰）方式下，草地生态系统小尺度上的空间属性进行分析后，得到以下研究结果： (1) 群落和物种的地上生物量以及土壤养分均表现出显著的小尺度空间结构特征，并且在不同利用方式影响下，这种空间属性表现出较大的差异。 (2) 禁牧（围封和刈割）草地的土壤有机碳和氮含量比较高。土壤有机碳的变程较小，变化范围为0.83～1.40 m，其在三种类型土地利用的变异系数都维持在18.0％左右;土壤全氮的空间自相关尺度为0.46～14.69m，变异系数为12.10％～30.86％。 (3) 群落地上生物量以禁牧（围封和刈割）草地较高。在围封禁牧和刈割禁牧草地内的变异系数分别为26.48％和19.94％，在自由放牧草地的变异系数为67.47％;其空间自相关尺度为0.74～3.9m。在不同的土地利用方式下，物种、功能群以及多样性指数都表现出显著的空间异质性，并且其空间属性变化各异。 (4) 土壤养分和植被的空间分布相互影响，互为因果关系。这种小尺度的空间格局差异是群落演替的驱动力，而不同的土地利用方式正是通过改变群落内小尺度的空间属性来影响了群落的演替方向。 在不同土地利用方式影响下，草地生态系统不仅在宏观功能，如生态系统生产力、群落结构和土壤的碳氮储量等方面表现出较大的差异，而且在微观结构上，如土壤养分和物种的空间属性等方面也发生了明显了变化，并且宏观和微观上的变化相互促进相互影响，它们的相互作用在很大程度上决定了群落的演替方向和草地生态系统全球变化过程中的具体作用。

全球变化背景下科尔沁沙地生态系统C、N循环过程

工业革命以来，由于人口的快速增加和人类活动的强烈干扰（主要包括煤炭、石油等石化燃料的燃烧、化肥生产和使用）导致土地利用/覆被变化、大气CO2浓度升高、N沉降等一系列全球环境变化问题。有关陆地生态系统生物地球化学循环，尤其是陆地生态系统C、N循环及其耦合过程方面的研究成为全球变化科学研究领域的重要内容。 干旱/半干旱地区占地球陆地总面积的1/3。与湿润地区相比较，干旱/半干旱地区生态系统稳定性比较差，往往属于生态脆弱区。因此，全球变化对干旱/半干旱地区生态系统影响更加敏感。科尔沁沙地位于我国北方干旱/半干旱地区，是我国典型的农牧交错区和生态脆弱区。科尔沁沙地是世界上人口密度最高的干旱/半干旱地区之一，人类活动对其影响剧烈。然而，有关科尔沁沙地生态系统C、N元素生物地球化学循环过程对土地利用/覆被变化、N沉降等全球变化响应及其反馈机制的研究非常缺乏。因此，本文以科尔沁沙地退化沙质草地、农田、不同年龄樟子松和杨树人工林等生态系统为对象，开展了造林、模拟N沉降和凋落物管理对生态系统C、N元素循环过程影响的研究。 在科尔沁沙地东南缘，以退化沙质草地、樟子松（Pinus sylvestris var. mongolica）人工林（15、24和30年生）、杨树（Poplus xiaozhuanica）人工林（7、11和15年生）为对象，研究草地转变为林地对生态系统C、N储量影响；以退化草地、榆树疏林草地和32年生樟子松人工林为对象，比较草地造林对土壤C、N循环过程及其土壤微生物性状的影响；以农田和5、10、15年生杨树人工林为对象，研究退耕还林对生态系统C、N储量和循环过程影响；以35年生樟子松人工林为对象，模拟研究N沉降和凋落物管理对生态系统C、N循环过程影响。通过上述研究，得到以下主要结果： （1）草地生态系统总C储量为34.38 Mg ha-1，15、24和30年生樟子松人工林生态系统总C储量分别为43.56、60.45和66.59 Mg ha-1，7、11和15年生杨树人工林生态系统总C储量分别为34.54、48.26和78.77 Mg ha-1；与农田相比，退耕5年的杨树人工林生态系统总C库储量下降13%，而10年和15年杨树人工林分别增加了176%和5倍；随着人工林年龄的增加，地上植被生物量C库储量占生态系统总C库储量的比例逐渐增加，并主要分配在树干。草地生态系统总N库储量为2.54 Mg ha-1，15、24和30年生樟子松人工林生态系统总N库储量分别为1.96、2.10和2.19 Mg ha-1，7、11和15年生杨树人工林生态系统总N库储量分别为2.27、1.84和2.60 Mg ha-1；与农田相比，退耕5年的杨树人工林生态系统总N库储量下降32%，而10年和15年杨树人工林分别增加了47%和76%；农田和草地造林后生态系统N储量依然主要分配在土壤中。 （2）草地和农田造林后土壤C、N库储量的变化受多因子的影响，例如林龄、树种种类以及立地条件等。农田和草地造林初期，土壤C、N库储量表现出下降趋势，随着林龄的增加，土壤C、N储量逐渐恢复。草地营造樟子松人工林30年后，0–60 cm深度土壤C、N储量依然显著低于草地；与草地相比，15年生杨树人工林土壤C、N储量差异不显著。在立地条件较好的情况下，10年杨树人工林土壤C、N储量已显著高于农田；然而，在立地条件相对较差的情况下，15年杨树人工林土壤C、N储量仍然与农田相比差异不显著。 （3）土地利用变化能够强烈地改变土壤C、N循环过程。与草地或疏林草地相比，32年生樟子松人工林土壤C、N、P含量显著降低；土壤C、N矿化过程发生显著变化，并且受季节变化的影响；在不同季节，土壤微生物量碳含量、代谢熵（qCO2）、微生物熵（MBC/TOC）以及土壤酶活性等在不同土地利用条件下表现出规律不一致。同样，农田退耕杨树人工林能够显著影响土壤C、N矿化过程，土壤无机氮（铵态氮+硝态氮）含量，土壤微生物量碳含量以及土壤微生物活性。草地造林在一定程度上导致土壤质量下降。而农田造林有利于土壤质量改善，尤其在在立地条件较好情况下。 （4）N添加增加对沙地樟子松人工林地上和地下C、N元素含量影响不大；N添加1年后，仅林下植被C、N含量显著增加，高氮处理（N15）凋落物N含量显著增加。N添加抑制了沙地樟子松人工林凋落物的早期分解和N、P元素释放。5、6、8和9月份土壤无机N含量均随着N输入增加表现出一定程度的增加，然而，7月份N添加导致土壤无机N含量降低。N添加对土壤潜在N矿化速率影响不显著。7和8月份N添加影响土壤C矿化速率，而其它月份影响不显著。低氮处理（N5）有利于增加土壤微生物量碳含量，而高氮处理（N15）在一定程度上降低土壤微生物量碳含量。 （5）凋落物输入变化（凋落物添加和凋落物移出）在一定程度上改变了35年生沙地樟子松人工林生态系统C、N循环过程。凋落物移出（C0）增加了林下植被C含量，降低了树木叶片N含量。凋落物移出抑制了凋落物分解和P元素的释放，而增加了C元素的早期释放速率，对N元素释放过程影响不显著。凋落物输入变化对不同月份土壤无机N含量和土壤N矿化过程影响均不显著。仅在6月份凋落物移出显著抑制了土壤C矿化速率，其它月份差异均不显著。凋落物管理对土壤微生物量碳含量影响不显著。 以上研究结果表明，土地利用变化、N沉降和凋落物输入改变等能够影响半干旱地区沙地生态系统C、N储量和循环过程。尤其是土地利用变化强烈改变沙地生态系统C、N储量、分配格局和循环过程，并且受到多因子的影响。科尔沁沙地樟子松人工林生态系统C、N元素生物地球化学循环存在密切的耦合关系。今后有必要进一步结合3S技术、同位素技术、模型模拟以及分子生物学技术等，从微观-宏观不同尺度上，研究半干旱地区沙地生态系统C、N循环过程对全球变化的响应及其反馈机制。

花椒主效化感物质分离鉴定及化感作用研究

近年来，随着对作物重茬障碍原因的深入研究，植物的化感作用越来越受到国内外众多学者的重视。花椒（Zanthoxy piperitum.）为芸香科植物，是一种收益早、用途广、价值高的经济树种，是川西干旱河谷地区的重要经济作物，其连作障碍也倍受关注，系统研究花椒化感作用将有助于理解和最终解决花椒连作障碍问题。本文首先通过萃取、层析等方法分离花椒主效化感成分；通过外加不同浓度的花椒叶水浸液研究了对土壤氮素养分循环的影响；研究了花椒叶水浸液对苜蓿生理生化、光合作用、氮素养分吸收的影响，并对外施氮肥对这种化感影响的缓解作用做了研究；研究了花椒化感潜力对全球变化——UV-B增强辐射的响应。主要研究结果如下： 1．用不同极性的有机溶剂对花椒叶水浸液浓缩浸膏萃取、柱层析，结合生物活性检测，分离得到主效化感作用组分的一种化感物质——对甲氧基苯酚。采用该物质纯品进行生物活性检测，证明其具有化感作用。 2．花椒叶水浸液处理土壤30天后，土壤硝态氮、铵态氮、无机氮（硝态氮＋铵态氮）与对照相比，随着花椒叶水浸液浓度的增加呈现降低的趋势，其中土壤铵态氮含量显著降低，而硝态氮含量的变化则不显著，无机氮含量也显著降低。土壤脲酶和蛋白酶的活性与无机氮含量的变化趋势相同。随着花椒叶水浸液浓度的增加，氨化细菌数量显著降低，固氮菌的数量变化不显著，硝化细菌和反硝化细菌数量有减少的趋势。60天后，硝态氮含量、铵态氮含量、无机氮随水浸液浓度增加的变化趋势与30天时相似；随着花椒叶水浸液浓度的增加，氨化细菌、固氮菌的数量显著减少，硝化细菌数量、反硝化细菌数量仍呈减少趋势；土壤脲酶、蛋白酶活性与第30天的变化趋势相同。第60天与第30天的结果相比，相同水浸液浓度处理的硝态氮、铵态氮、无机氮均有下降的趋势，但除了25g.L－1水浸液处理的外，其它相同浓度的处理间差异均不显著；除了12.5 g.L的处理外土壤脲酶活性均呈增强的趋势；蛋白酶活性都有不同程度的增加；花椒叶水浸液处理的土壤硝化细菌和反硝化细菌数量呈增加趋势。 3．随着花椒叶水浸液浓度的增加，显著抑制了苜蓿根长、地上地下生物量、叶绿素含量、叶片中可溶性蛋白的含量，净光合速率。苜蓿体内四种抗氧化酶（POD、SOD、CAT、APX） 活性随着水浸液浓度的增加而降低，而丙二醛含量则增加。苜蓿氮初级同化相关酶硝酸还原酶（NR）、谷氨酰合成酶(GS)、谷氨酸脱氢酶（GDH）的活性随着水浸液浓度的增加受到不同程度的影响。总的来说，苜蓿硝酸还原酶、谷氨酰合成酶的活性受到抑制，而谷氨酸脱氢酶活性的变化则比较复杂，根呈先降低后增加的趋势，叶片则无显著变化。外施两种不同浓度的硝酸铵氮肥后，对12.5、25 g.L－1花椒叶水浸液处理的苜蓿化感作用有显著的缓解作用，表现在株高、生物量、光合作用等方面，大多达到与对照（0 g.L－1）未施氮肥无显著差异的水平，而对50 g.L－1水浸液处理的苜蓿幼苗，虽有一定的缓解作用，但这种作用均未达到与对照（0 g.L－1）未施氮肥时无显著差异的水平。 4． UV-B增强辐射处理花椒后，花椒的化感潜力显著增强。花椒叶片内UV-B吸收物质的含量和总酚含量均显著增加。 In recent years, with profound research on the reasons of continuous cropping obstacles, allelopathy received increasing attention to many scholars at home and abroad. Zanthoxy bungeanum as a Rutaceae plant is a high economic value species which gains early and uses widely. Zanthoxylum is an important economic crop in the arid valley of western Sichuan region, and its not even has received much concern for the continuous cropping obstacles. The systematic study of allelopathy of Zanthoxylum will contribute to the understanding and final settlement of this issue. The major allelopathic composition was separated through the extraction, chromatography combined with other methods. The impact on soil nutrient cycling was also studied through the addition of different concentrations of water extracts of Zanthoxylum. Furthermore, the effects of water extracts of Zanthoxylum leaves on alfalfa leaf physiological and biochemical indexes, photosynthesis, soil enzymes and nutrient uptake of nitrogen and the mitigation of allelopathy through using external fertilizer were studied to put forward scientific resolvent for Zanthoxylum continuous cropping obstacles .The response of allelopathic potential of Zanthoxylum to global change - UV-B enhanced radiation was studied . The main findings are as follows: 1. Through extraction with different polar organic solvents on concentrated water extract of Zanthoxylum leaf and then using column chromatography combined with detection of biological activity, one of the main allelopathic components- methoxy-phenol was isolated. The biological activity testing of the pure material of methoxy-phenol proved that it does have allelopathic potential. 2. Thirty days after treating soil with water extract of Zanthoxylum leaf, as compared with the control, the contents of soil nitrate, ammonium, nitrate plus ammonium nitrogen showed a trend of decrease with the increase of the concentration of water extract whereas the content of ammonium nitrogen showed a significant reduction, and the content of nitrate did not change significantly, the content of nitrate plus ammonium nitrogen also showed a significant (P <0.05) redction. The activity of soil urease and protease showed the same trend as the content of nitrate nitrogen plus ammonium nitrogen. With the increase in the concentration of water extract, the number of ammonification bacteria significantly reduced but nitrogen-fixing bacteria did not change significantly and there was a decreasing trend in the number of nitrifying bacteria and denitrifying bacteria. Sixty days after the treatment, with the increase in solution concentration of water extract of Zanthoxylum leaf, the content of nitrate、 ammonium nitrogen, nitrate plus ammonium nitrogen showed a similar change trend to 30 days’; the number of ammonification bacteria, nitrogen-fixing bacteria significantly reduced ; the number of nitrifying bacteria, denitrifying bacteria was still an downward trend; the activity of soil urease and protease showed the same trend as the 30th days’. Compared to the results of the 30th days’, the content of nitrate, ammonium, nitrate plus ammonium nitrogen showed a decrease trend between the treatment of same concentration, but there was no significant difference except the treatment of 25g.L-1 between the same concentration; the activity of soil urease showed enhanced trend except the treatment of 12.5 g.L-1; the activity of protease increased to varying degrees; the number of ammonification bacteria、 nitrifying bacteria and denitrifying bacteria were growing while nitrogen-fixing bacteria reduced.. 3. With the increase of the concentration of water extract of Zanthoxylum leaf, the water extract significantly inhibited the root length, aboveground biomass, content of chlorophyll and soluble protein in leaf and net photosynthetic rate. The activity of four antioxidant enzymes (POD, SOD, CAT, APX) reduced with the increase in concentration of the water extract but the content of MDA increased. The activity of enzymes related to primary nitrogen assimilation such nitrate reductase (NR), glutamyl synthetase (GS), glutamate dehydrogenase (GDH) were subject to different degrees with an increase in the concentration of water extracts. In general, the activity of nitrate reductase, glutamyl synthetase were inhibited, while change in the activity of glutamate dehydrogenase was more complex. The activity of glutamate dehydrogenase in leaf was first reduced and then increase,but did not change significantly in root. After using two external different concentrations of nitrogen fertilizer, there was a significant mitigation in inhibiton in plant height, biomass, photosynthesis, etc. in the treatment of 12.5,25 gL-1 of water extract of Zanthoxylum leaf, and most of these indexes showed no significant difference with the control (0 g.L-1, no external fertilizer was added) .Although there showed a certain degree of ease in the treatment of 50 g.L-1 , there was still a significant difference compared with the control (0 gL-1) in which no external fertilizer was used. 4.The allelopathic potential of Zanthoxylum positively responded to enhanced UV-B significantly. The content of UV-B absorbing compounds and the total phenol also significant increased.

水分亏缺和凋落物水浸液对云杉幼苗生长和种子萌发的影响

在人类活动导致全球变暖的前提下，由于全球气温的升高，地表水分加速向空中蒸发。从20世纪70年代至今，地球上严重干旱地区的面积几乎扩大了一倍。这一增长的一半可归因于气温升高而不是降雨量下降，因为实际上同期全球平均降水量还略有增长。干旱对陆地植物和农林生态系统产生深远影响，并已成为全球变化研究的一个重要方面。位于青藏高原东部的川西亚高山针叶林是研究气候变暖对陆地生态系统影响的重要森林类型。森林采伐迹地、人工林下和林窗环境作为目前该区人工造林和森林更新的重要生境，其截然不同的光环境对亚高山针叶林更新和森林动态有非常重要的影响。凋落物产生的化感物质可通过影响种子萌发和早期幼苗的定居而影响种群的建立和更新，而人工林和自然林物种以及更新速度的差异性也都受凋落物的影响。 云杉是川西亚高山针叶林群落的重要树种之一,在维持亚高山森林的景观格局和区域生态安全方面具有十分重要的作用，其自然更新能力及其影响机制一直是研究的热点问题。本试验以云杉种子和2年生幼苗为研究对象，从萌发、根尖形态、幼苗生长、光合作用、渗透调节和抗氧化能力等方面研究了不同光环境下水分亏缺和凋落物水浸液对云杉种子和幼苗生长的影响。旨在从更新的角度探讨亚高山针叶林自然更新的过程，其研究成果可在一定程度上为川西亚高山针叶林更新提供科学依据，同时也可为林业生产管理提供科学指导。主要研究结论如下： 水分亏缺在生长形态、光合作用、抗氧化能力、活性氧化对云杉幼苗都有显著影响。总体表现为，水分亏缺导致了云杉幼苗的高度、地径、单株总生物量降低，增加了地下部分的生长；水分亏缺显著降低了云杉叶片中相对含水量、光合色素、叶氮含量，净光合速率和最大量子产量(Fv/Fm)，提高了幼苗叶片中膜脂过氧化产物（MDA）的含量；水分亏缺提高了幼苗叶片中过氧化氢(H2O2)含量，超氧荫离子（O2-）生成速率以及脯氨酸和抗氧化系统的活性（ASA, SOD, CAT, POD, APX和GR）。从这些结果可知，植物在遭受水分亏缺导致的伤害时，其自身会形成防御策略，并通过改变形态和生理方面的特性以减轻害。但是，这种自我保护机制依然不能抵抗严重水分亏缺对植物的伤害。 模拟林下低光照条件显著增加单株植物的地上部分生长，尤其是其叶片的比叶面积（叶面积/叶干重），同时其光合色素含量和叶片相对含水量也显著增加，这些改变直接导致植株光合速率和生物量的增加。同时，与高光照水平相比，低光照幼苗的膜脂过氧化产物（MDA）和活性氧物质均较低，显示出低光照比高光照水平对植物的更低的氧化伤害。尽管低光照也导致大部分抗氧化酶活性降低，但这正显示出植物遭受低的氧化伤害，更印证了前面的结论。 凋落物水浸液影响了云杉种子的萌发和根系的生长，更在形态、光合作用、抗氧化能力、活性氧物质以及叶氮水平上显著影响了云杉幼苗，其中，以人工纯林凋落物的影响更有强烈。具体表现在，种子萌发速率和萌发种子幼根的长度表现为对照>自然林处理>人工纯林；凋落物水浸液抑制种子分生区和伸长区的生长，人工林处理更降低了根毛区的生长，使根吸水分和养分困难。对2年生幼苗的影响主要表现在叶绿素含量、光合速率以及叶氮含量的降低；膜脂过氧化产物、活性氧物质和抗氧化酶系统的显著增加。同样的，人工纯林处理对云杉幼苗的影响显著于自然林处理。 在自然生态系统中，由于全球变暖气温升高导致的水分亏缺和森林凋落物都存在森林的砍伐迹地，林窗和林下环境中。我们的研究表明，与迹地或林窗强光照比较，林下的低光照环境由于为植物的生长营造了较为湿润的微环境，因此水分亏缺在林下对云杉幼苗造成的影响微弱。这可以从植物的形态、光合速率以及生物量积累，过氧化伤害和抗氧化酶系统表现出来。另一方面，凋落物水浸液在模拟林下低光照环境对植物的伤害也微弱于强光照环境，这与强光照环境高的水分散失导致环境水分亏缺有关；而人工纯林处理对云杉幼苗的伤害比对照和自然林处理显示出强烈的抑制作用。 Under the pre-condition of global warming resulted from intensive human activities, water in the earth’s surface rapidly evaporates due to the increase of global air temperature. From 1970s up to now, the area of serious drought in the world is almost twice as ever. This increase might be due to the increasing air temperature and not decreasing rainfall because global average rainfall in the corresponding period slightly is incremental. Drought will have profound impacts on terrestrial and agriculture-forest system and has also become the important issue of global change research. The subalpine coniferous forests in the eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying the effects of global warming on terrestrial ecosystems. The light environment significantly differs among cutting blanks, forest gap and understory, which is particularly important for plant regeneration and forest dynamics in the subalpine coniferous forests. Picea asperata is one of the keystone species of subalpine coniferouis forests in western China, and it is very important in preserving landscape structure and regional ecological security of subalpine forests. The natural regeneration capacities and influence mechanism of Picea asperata are always the hot topics. In the present study, the short-term effects of two light levels (100% of full sunlight and 15% of full sunlight), two watering regimes (100% of field capacity and 30% of field capacity), two litter aqueous extracts (primitive forest and plantation aqueous extracts) on the seed germination, early growth and physiological traits of Picea asperata were determined in the laboratory and natural greenhouse. The present study was undertaken so as to give a better understanding of the regeneration progress affected by water deficit, low light and litter aqueous extracts. Our results could provide insights into the effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientific direction for the forest production and management. Water deficit had significant effects on growth, morphological, physiological and biochemical traits of Picea asperata seedlings. Water deficit resulted in the decrease in height, basal diameter, total biomass and increase in under-ground development; water deficit significantly reduced the needle relative water content, photosynthetic pigments, needle nitrogen concentration, net photosynthetic rate and the maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA) in Picea asperata seedlings; water deficit also increased the rate of superoxide radical (O2-) production, hydrogen peroxide (H2O2) content, free proline content and the activities of antioxidant systems (ASA, SOD, POD, CAT, APX and GR) in Picea asperata seedlings. These results indicated that some protective mechanism was formed when plants suffered from drought stress, but the protection could not counteract the harm resulting from the serious drought stress on them. Low light in the understory significantly increased seedling above-ground development, especially the species leaf area (SLA), and photosynthetic pigments and relative needle content. These changes resulted in the increase in net photosynthetic rate and total biomass. Moreover, the lower MDA content and active oxygen species (AOS) (H2O2 and O2-) in low light seedlings suggested that low light had weaker oxidative damage as compared to high light. Lower antioxidant enzymes activities in low light seedlings indicated the weaker oxidative damage on Picea asperata seedlings than high light seedlings, which was correlative with the changes in MDA and AOS. Litter aqueous extracts affected seed germination and root system of Picea asperata seedlings. Significant changes in growth, photosynthesis, antioxidant activities, active oxygen species and leaf nitrogen concentration were also found in Picea asperata seedlings, and plantation treatment showed the stronger effects on these traits than those in control and primitive forest treatment. The present results indicated that seed germination and radicle length parameters in control were superior to those in primitive forest treatment, and those of primitive forest treatment were superior to plantation treatment; litter aqueous extracts inhibited the meristematic and elongation zone, and plantation treatment caused a decrease in root hairs so as to be difficult in absorbing water and nutrient in root system. On the other hand, litter aqueous extracts significantly decreased chlorophyll content, net photosynthetic rate and leaf nitrogen concentration of Picea asperata seedlings; MDA, AOS and antioxidant system activities were significantly increased in Picea asperata seedlings. Similarly, plantation treatment had more significant effect on Picea asperata seedlings as compared to primitive forest treatment. In the nature ecosystem, water deficit resulted from elevating air temperature and litter aqueous extract may probably coexist in the cutting blank, forest gap and understory. Our present study showed that water deficit had weaker effects on low light seedlings in the understory as compared to high light seedlings in the cutting blank and forest gap. The fact was confirmed from seedlings growth, gas exchange and biomass accumulation, peroxidation and antioxidant systems. This might be due to that low light-reduced leaf and air temperatures, vapour-pressure deficit, and the oxidative stresses can aggravate the impact of drought under higher light. On the other hand, litter aqueous extracts in the low light had weaker effects on the Picea asperata seedlings than those at high light level, which might be correlative to the water evapotranspiration under high light. Moreover, plantation litter aqueous extracts showed stronger inhibition for seed germination and seedling growth than control and primitive forest treatments.

增温对川西北高寒草甸植物群落结构及几种主要植物生长和生理的影响

由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20世纪升高了0.6 ℃，并预测在本世纪将上升1.4-5.8 ℃。气候变暖对陆地植物和生态系统影响深远，并已成为全球变化研究的重要议题。高海拔、高纬度地带的生态系统对气候变化最敏感。而在高原和高山极端环境影响下所形成的高寒草甸生态系统极其脆弱，对由于温室效应引起的全球气候变化极其敏感，对这些变化的响应更具有超前性。 本研究以川西北高寒草甸植物群落及几种主要物种为研究对象，采用国际山地综合研究中心（ITEX）普遍所采用的增温方法-----开顶式生长室（OTC）模拟气候变暖来研究增温对高寒草甸植物群落结构、物质分配及其主要物种生长和生理的影响，以探讨高寒草甸植物响应与适应气候变暖的生物学和生态学机制。主要研究结论如下： 1、OTC的增温效果 由于地温、地表温度和气温的平均值在OTC内分别高出对照样地0.28℃、0.46℃和1.4℃，这说明本研究所采用的开顶式生长室（OTC）起到了增温的作用；同时，由于温室内与温室外接受的降水量相同，温室内由于热量条件的改善，土壤蒸发和植被的蒸腾作用增强，直接导致了OTC内土壤表层相对湿度的减少。 2、群落结构对增温的响应 由于增温时间较短，增温内外样地的物种组成并未发生改变；但增温后一定程度上改变了植物群落的小气候环境，从而导致物种间的竞争关系被破坏，种间竞争关系的破坏引起群落优势种组成发生相应的改变，在对照样地，鹅绒委陵菜、甘青老鹳草、遏蓝菜和蚤缀是占绝对优势的物种，而在OTC内，小米草、尼泊尔酸模、垂穗披碱草、发草和羊茅的重要性显著增加。 禾草和杂草由于对增温的生物学特性及其资源利用响应的不同，加之增温造成土壤含水量下降等环境因子的改变。与对照样地相比较，OTC内禾草的盖度及生物量都显著增加，而杂草的盖度和生物量则显著下降。 3、植物生长期对增温的响应 OTC内立枯和调落物的生物量在生长季末（10月份）都要小于对照样地的立枯和调落物生物量，而OTC内的地上鲜体生物量在10月份却略高于对照样地。这说明OTC内植物的衰老或死亡得以延缓，而植物的生长期得以延长。 4、群落生物量及分配对增温的响应 OTC内的地上鲜体生物量(10月份除外)和地下0-30cm的根系生物量与对照样地相比较，都出现了不同程度的减少；土壤根系的分配格局也发生了明显的改变，其中，OTC内0-10cm土层的生物量分配比例增加，而20-30cm土层生物量分配比例的减少。 5、群落碳、氮对增温的响应 增温后，OTC内植物群落地上活体和地下活根的碳浓度不同程度的高于对照样地，植物群落的碳库在OTC内也略高于对照样地；而OTC内植物群落地上活体和地下活根的氮浓度不同程度的低于对照样地，其植物群落的氮库与对照样地相比也略有下降。 6、几种主要植物的生长及物质分配对增温的响应 垂穗披碱草在增温后株高、比叶面积和地上生物量均显著地增加；尼泊尔酸模在增温后比叶面积和单株平均生物量积累显著地增加，而各组分中，增温处理使叶的生物量显著增加，而根的生物量却显著下降；鹅绒委陵菜在增温后株高、比叶面积和单株平均生物量积累显著地减少，而各组分中，增温处理使叶和茎的生物量显著减少，根的生物量却显著地增加。 尼泊尔酸模的LMR、RMR、R/S、根部碳含量、碳和氮在叶片与根部的分配比例在增温后显著地增加，而SMR、根部氮含量、碳和氮在茎部的分配比例在增温后却显著地降低；鹅绒委陵菜的RMR、R/S、碳和氮在根部的分配比例在增温后显著地增加，而SMR、LMR、碳在叶片的分配比例在增温后却显著地降低 7、几种主要植物的光合生理过程对增温的响应 增温使垂穗披碱草和尼泊尔酸模叶片中的叶绿素a、叶绿素b、总叶绿素含量显著增加；而鹅绒委陵菜叶片的叶绿素a、叶绿素b、总叶绿素含量在增温后显著减少，类胡萝卜素含量在增温后却显著增加。 增温对3种植物的气体交换产生了显著影响。其中，垂穗披碱草和尼泊尔酸模叶片的光响应曲线在增温后明显高于对照处理，A、E、gs、Pmax、、Rday、AQY和LSP显著增加，而LCP则显著降低；鹅绒委陵菜的光响应曲线在增温后则明显的低于对照处理，A、E、gs、Pmax、、Rday、AQY和LSP显著减少，而LCP则显著增加。 增温后垂穗披碱草和尼泊尔酸模叶片的Fv/Fm、Yield和qP显著增加；而鹅绒委陵菜叶片的Fv/Fm、Yield和qP则显著减少，qN却显著地增加。 8、几种主要植物的抗氧化酶系统对增温的响应 增温使垂穗披碱草和尼泊尔酸模体内抗氧化酶活性和非酶促作用有所提高，植物膜脂过氧化作用降低；鹅绒委陵菜叶片中酶促反应和非酶促反应在增温后也显著提高，但可能由于增温后的土壤干旱超过了鹅绒委陵菜叶的抗氧化保护能力，抗氧化酶活性及非酶促反应（脯氨酸、类胡萝卜素）的提高不足以完全清除干旱诱导形成的过量活性氧，因此叶片的膜脂过氧化程度仍然显著提高。 Enrichment of atmospheric greenhouse gases resulted from human activities such as fossil fuel burning and deforestation has increased global mean temperature by 0.6 ℃ in the 20th century and is predicted to increase in this century by 1.4-5.8 ℃. The global warming will have profound, long-term impacts on terrestrial plants and ecosystems. The ecoologcial consequences arising from global warming have also become the very important issuses of global change research. The terrestrial habitats of high-elevation and high-latitude ecosystems are regarded as the most sensitive to changing climate. The alpine meadow ecosystme, which resulted from the composite effects of mountain extreme climatic factors in Tibetan Plateau, is thus thought to be especially vulnerable and sensitive to global warming. In this paper, the response of plant community and several main species in the alpine meadow of Northewst Sichuan to experimemtal warming was studied by using open-top chambers (OTC). The aim of the this study was to research the warming effects on plant community structure, substance allocation, growth and physiological processes of several mian species, and to explore the biological and ecological mechanism of how the alpine meadow plants acclimate and adapt to future global warming. The results were as follows: 1. Warming effects of OTC The mean soil temperature, soil surface temperature and air temperature in OTC manipulation increased by 0.28℃、0.46℃ and 1.4℃ compared to the control during the growing season. This suggested that the OTC used in our study had increased temperature there. Meanwhile, the OTC manipulation slightly altered thermal conditions, but the same amount of precipitation was supplied to both the OTC manipulation and the control, so higher soil evaporation and plant transpiration in OTC manipulation directly lead to the decrease of soil surface water content. 2. The reponse of community structure to experimental warming The species richness was not changed by the short-term effect of OTC manipulation. However, experimental warming changed the microenvironment of plant community, therefore competitive balances among species were shift, leading to changes in species dominance. In the present study, the dominant plant species in the control plots were some forbs including Potentilla anserine, Geranium pylzowianum, Thlaspi arvense and Arenaria serpyllifolia, however, the importance value of some gramineous grasses including Elymus nutans, Deschampsia caespitosa, Festuca ovina, and some forbs including Euphrasia tatarica and Rumex acetosa significantly increased in OTC. The different biology characteristics and resource utilizations between gramineous grasses and forbs, and enhanced temperature caused change in some environment factors such as soil water content. As a result, the coverage and biomass of gramineous grasses significantly increased in OTC compared to the control, however, the coverage and biomass of forbs singnifciantly decreased in OTC compared to the control. 3. The reponse of plant growing season to experimental warming Both the standing dead and fallen litter biomass in OTC were lower than those in the control in October, and the biomass of aboveground live-vegetation in OTC was higher than that of the control. The results indicated that the senescence of plants was postponed, and the growing season was prolonged in our research. 4. The reponse of community biomass accumulation and its allocation to experimental warming Experimental warming caused the decrease of aboveground live biomass and belowground root biomass except for the aboveground live biomass in October. Experimental warming also had pronounced effects on the pattern of root biomass allocation. In the present study, the root biomass in 0-10cm soil layer increased in OTC manipulation compared to the control, however, the root biomass in the 20-30cm soil layer decreased in OTC manipulation compared to the control. 5. The reponse of community C and N content to experimental warming The C concentration and stock in aboveground live and belowground root both increased in OTC manipulation compared to the control. However, the N concentration and stock in aboveground live and belowground root both decreased in OTC manipulation compared to the control. 6. The reponse of gowth and biomass, C and N alloction of several species to experimental warming Experimental warming significantly increased the height, SLA (specific leaf area) and aboveground biomass of Elymus nutans in OTC manipulation compared to the control. The SLA and total biomass of Rumex acetosa also significantly increased in OTC manipulation compared to control, among the different components of Rumex acetosa, leaf biomass significantly increased, but root biomass significantly decreased in OTC manipulation compared to the control. However, the height, SLA and total biomass of Potentilla anserina significantly decreased in OTC manipulation compared to the control, among the different component of Potentilla anserina, leaf and stem biomass significantly decreased, but root biomass significantly increased in OTC manipulation compared to the control. The LMR (leaf mass ratio), RMR (root mass ratio), R/S (shoot/root biomass ration) and root C concentration of Rumex acetosa significantly increased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively more C and N content to leaf and root in response to experimental warming, however, the SMR (stem mass ration) and root N concentration of Rumex acetosa significantly decreased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively less C and N content to stem in response to experimental warming. The RMR and R/S of Potentilla anserina significantly increased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively more C and N content to root in response to experimental warming, however, the SMR and LMR of Potentilla anserina significantly decreased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively less C and N content to leaf in response to experimental warming. 7. The reponse of physiological processes of several species to experimental warming Experimental warming significantly increased chlorophyll a, chlorophyll b and total chlorophyll of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control. However, chlorophyll a, chlorophyll b, total chlorophyll and carotenoid of Potentilla anserina in OTC manipulation significantly decreased compared to outside control. Experimental warming had pronounced effects on gas exchange of Elymus nutans, Rumex acetosa and Potentilla anserine. In the present study, warming markedly increased the light response curves of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control, and also singnificantly increased A (net photosynthesis rate), E (transpiration rate), gs (stomatal conductance), Pmax (maximum net photosynthetic rate), Rday (dark respiration rate), AQY (apparent quantum yield) and LSP (light saturation point), but LCP (photosynthetic light compensation) of Elymus nutans and Rumex acetosa in OTC manipulation singnificantly decreased compared to outside control. However, warming markedly decreased the light response curves of Potentilla anserina in OTC manipulation compared to outside control, and also singnificantly decreased A, E, gs, Pmax, Rday, AQY and LSP, but LCP of Potentilla anserina in OTC manipulation singnificantly increased compared to outside control. Experimental warming singnificantly increased the chlorophyll fluorescence kinetics parameters such as Fv/Fm, Yield and qP of Elymus nutans and Rumex acetosa and qN of Potentilla anserina in OTC manipulation, but Fv/Fm, Yield and qP of Potentilla anserina in OTC manipulation singnificantly decreased. 8. The reponse of antioxidative systems of several species to experimental warming Experimental warming tended to increase the activities of antioxidative enzymes and stimulate the role of non-enzymes of Elymus nutans and Rumex acetosa. As a result, MDA content of Elymus nutans and Rumex acetosa decreased. The activities of antioxidative enzymes and non-enzymes of Potentilla anserina also significantly increased in OTC manipulation, but more O2- was produced because of lower soil water content, and the O2- accumulation exceeded the defense ability of antioxidative systems and non-enzymes fuctions. As a result, MDA content of Potentilla anserine still increased in OTC manipulation compared to outside control.

基于RS和GIS的四川省土地利用/覆盖变化及可持续利用研究

LUCC是全球变化研究的核心主题之一，也是社会经济可持续发展的关键问题。改革开放后四川的社会经济发展非常快，在各种因素的驱动下，土地利用/覆盖发生了深刻变化。目前四川省缺乏基于实际调查数据的、全域性的、具有连续时间序列的LUCC和驱动力分析及土地可持续利用研究成果，这对我们从全局上把握全省土地利用现状、发展变化趋势，利用土地政策参与宏观调控，实现长期可持续发展目标，建设资源节约型、环境友好型社会极为不利。本研究针对这一问题，选取全川八大土地利用类型作为研究对象，研究了全省1996年到2006年的土地利用/覆盖格局和变化情况，分析了不同尺度的驱动因素，对全省农用地和建设用地的集约利用状况、潜力进行了分析评价，并提出相应的对策措施。 1．1996年-2006年10年来整个省域的土地利用/覆盖格局变化。 （1）1996年-2006年全省的土地利用/覆盖格局 1996年，全省是一个以农用地为主的土地利用/覆盖格局，林地和牧草地属于优势覆盖类型（合占69.17%），居民点及工矿用地和交通用地合占只有3%左右。 2000年的LUCC格局较为明显的特点是耕地所占比重下降0.4个百分点，水域和未利用土地所占比重有所下降，牧草地保持不变，其余地类所占比重有所上升。 与2000年相比，2004年林草地的优势格局进一步得到强化（合占比重达到70.23%）。耕地面积占幅员面积的比重下降0.83个百分点，略有下降的有未利用土地、水域和牧草地。值得关注的是在“退耕还林还草”的大背景下，牧草地占幅员面积的比重下降0.04个百分点。 到2006年，仍为林草地为主导优势的格局，二者合占上升0.15%。在城市化快速推进的背景下，居民点及工矿用地中的城市用地和建制镇用地占比重超过15%，农村居民点占比重降至76%。交通用地中农村道路占比重降至57.8%,公路用地占比重升至37.5%。五个地貌区的土地利用/覆盖格局与全省的变化基本一致。值得关注的是盆西平原区的交通用地上升幅度和盆地丘陵区的未利用土地的开发利用力度明显大于其它地貌区。 （2）1996-2006年10年间土地利用/覆盖格局的变化 1996-2000年4年间，耕地、水域和未利用地三个地类下降，年均减少0.75、0.19和0.32个百分点。其中耕地年均减少49229.0公顷，约一半流向林地，13.77%流向园地，约20%流向建设用地。另外5个地类面积增长，增长绝对量最大的是林地，年均增长40063.7公顷，交通用地增幅最大，4年年均增长达1.95%。 2001-2004年是西部大开发逐步推进、“退耕还林还草”项目全面展开和土地整理深入实施的关键期，LUCC更为深刻。耕地、未利用地、水域和牧草地四个地类面积下降，其余地类按增长幅度依次是园地、交通用地、居民点及工矿用地和林地。耕地加速下降，年均降幅达到1.59%，其减少去向主要是林地（占66.75%）和园地（占19.84%），其增加来源主要是未利用地、园地和水域。交通用地的增幅最大，为3.96%,其增加主要来源于耕地、未利用土地和林地，分别占49.96%、16.63%和13.09%。居民点及工矿用地增长幅度为3.12%。 从1996年到2006年的10年间，耕地、未利用地、水域和牧草地下降幅度分别为10.36%、3.61%、1.34%和0.26%。园地增幅达23.61%。绝对面积增长最大的则是林地，达630733.3公顷。交通用地和居民点及工矿用地增幅也较大，分别为15.00%和9.31%。 10年间年均总变化量为310326.6公顷，2000年-2004年之间变化最大（为356865.8公顷），高于平均变化量，而1996-2000年间和2004-2006年间都小于平均变化量。 （3）10年间不同地貌区的LUCC变化 盆西平原区的特点是园地大幅上升达77%，居民点及工矿用地和交通用地也大幅上升，耕地、未利用地下降幅度大，该区耕地、水域、未利用地的减少强度和园地、居民点及工矿用地、交通用地的上升强度均居五区第一；盆地丘陵区的特点是牧草地下降幅度大，为-36.89%,交通用地、园地和林地上升幅度较大，该区耕地减少、未利用地减少、林地增加、居民点及工矿用地和交通用地增加的变化强度均居五区相应地类增减的第二位；盆周山地区的特点是耕地减少较多，交通用地和园地增长较大，该区林地变化强度居各区第一位，牧草地和水域变化强度居各区第二位，耕地、居民点及工矿用地和未利用地居各区第三位；川西南山地区的特点是园地、耕地、交通用地和居民点及工矿用地变化幅度大，另外四个地类变化较小。该区减少的牧草地占全省牧草地减少的97.91%，变化强度居各个地貌区的第一位，园地相对变化强度居五区的第二位；川西北高山高原区的特点是耕地大幅下降、园地大幅上升，交通用地升幅也较大，其余地类变化不大。值得注意的是，该区牧草地和水域面积增加，与全省该地类的变化相反。其余地类的相对变化强度均是五个地貌区中最小的。 用变化强度分值考量变化强度，盆西平原区的变化强度最大，盆地丘陵区和盆周山地区的变化强度相当，川西北高山高原区的变化强度则要小得多。 （4）1996年及2006年全省土地利用/覆盖格局的景观生态学分析 全省是以自然景观占优势（占约70%）、农业景观为补充、建设用地景观居于从属地位的土地利用景观格局。景观多样性和均匀度不高。到2006年，全省总的景观格局并无大的改变。总体情况是随着时间的推移和人类活动的加强，区内景观优势度上升、多样性和均匀度变小。但斑块数减少，斑块面积和斑块孔隙度有所增大。斑块的形状指数和分维数均有所下降，表明受人为干扰有加剧的趋势。反映景观格局结构的破碎度指数有轻微下降。景观指数的变化表明全省土地利用有缓慢集中、规模聚集的趋势。 （5）三大生态建设工程对土地利用/覆盖变化的影响 1996-2006年间LUCC与三大生态建设工程实施的耦合分析，发现退耕工程对耕地、林地、牧草地等地类覆盖变化的影响最大，天保工程次之，长防工程最小。 2.四川省LUCC驱动力分析 （1）总体分析： 从整体上分析，人为因素对区域整体LUCC的影响从1996年的63.32%增加到2006年的66.99%，变得日益强烈。同时人为因素影响强度表现出明显的区域差异，地势平缓、经济区位条件好的区域其人为影响强度明显较高。 政策体制转变下的经济高速增长、快速的城市化、工业化过程和生态建设是四川省LUCC宏观尺度的驱动因素。区域的LUCC主要受到了由内向外(从城市到乡村)和由外向内(从山顶向平地)两种作用力的共同推动。局部尺度上，如距离交通线、水利线、中心城市的远近，地形凸起、大型独立项目落址、重污染项目的阻隔等，甚至一些乡规民俗等因素也会成为LUCC的驱动影响因素。在较小的尺度上,人类个体行为选择对LUCC的影响也是存在的。 　　根据驱动因子的特性作者将其划分为驱变、阻变、良性、惰性因子等类型。 （2）分地貌区的驱动因子分析 各地貌区都存在城市化、工业化、生态工程实施、自然灾害等驱动因子，但主次不一。对于盆西平原和盆地丘陵区，城市化、工业化是前两位的因子，而对另外三个地貌区，生态工程实施和产业结构调整则成为第第一、二位的驱动因子。 （3）分地类的驱动因子分析（以坡耕地为例） 分坡度的耕地变化分析发现，耕地减少主要集中在2°以下的平地、15°-25°和25°以上三个坡度级，是其它坡度级耕地减幅的三倍左右。这表明耕地减少受城市化进程和“退耕还林还草”工程驱动影响尤为巨大。 3.土地利用格局优化、集约利用评价和可持续利用及对策研究 （1）土地利用格局优化的战略选择及调整预测 土地利用格局调整的战略是农业生产用地、建设用地和生态及其他用地占幅员的比重分别稳定在13%、7%和80%左右，重点是三大类别内部二级和三级地类的合理调整。 （2）全省土地集约利用评价 全省农用地利用集约度为0.46，总体上集约度不高，处于较适度利用阶段。建设用地利用集约度为0.38，处于较适度利用阶段。集约利用提升空间较大。 农用地的潜力主要在于加强土地保育、完善利用制度、提高单产。城市建设用地的包括存量潜力、强度潜力、结构潜力，空间很大。农村居民点整理潜力可以逐步挖掘。 （3）新增建设用地集约利用的统筹安排 据测算，到2020年，四川省城市建设用地需求量在463850-492360hm2之间，城镇各业新增建设用地规模为361276.79hm2，占用耕地200565.94 hm2。2004-2020年间四川省农村居民点整理潜力33.86万hm2。农村居民点建设用地需求量为70.57万公顷。 （4）土地集约利用措施与坡耕地可持续利用战略 提出了土地集约利用的措施。在对坡耕地生态系统结构与功能分析的基础上，提出坡耕地可持续利用战略与生态恢复战略，并从技术和政策层面提出了坡耕地合理利用和生态退耕的措施和建议。 LUCC is one of the key questions of global change and sustainable development of society. After the opening and reform of China, the society and economy of Sichuan Province developed very fast ,the land-use/cover changed very strong droved by many factors .But nowadays we have no constant spatial-temporal study and driving force analysis about the whole province based on investigation. And it is lack of land sustainable utilization study based on correlative study. So we choose all the land resource in Sichuan, combine RS and GIS and field investigation, and take statistic-mathematic means and system analysis, to study the LUCC patterns and different scale driving force of different physiognomy regions, land cover types and periods; to analyze the current situation and potential of land resource intensive utilization, and gave out corresponding measurements. We found that forest and grassland are the dominant cover types of Sichuan provincial land –use/cover pattern, and becoming more and more stronger from 1996 to 2006,the natural landscape is the metric and occupy 70%,the diversity and evenness index are not high; the totally change quantity from 2000 to 2004 is the biggest; cultivated land especially steep cultivated land ,garden plot, forestry land ,settlement and industry land and traffic land changed relative stronger; among five physiognomy regions ,the changing intensity of PEN XI PING YUAN QU is the biggest, CHUAN XI BEI GAO SHAN GAO QU is smallest; under the background of policy system changing, the fast developing of economy, fast urbanization and industrialization and ecology construction are the macro-scale driving force of Sichuan provincial LUCC; to compare the impacts of “TUI GENG GONG CHENG” on LUCC especially to cultivated land ,forestry land and grassland is strongest, “TIAN BAO GONG CHENG ” is stronger,“ HANG FANG GONG CHENG” is smallest; the intensive utilization level of farmland and construction land of whole province is relative moderation, there is huge potential to excavate and fulfill the increasing demand of construction land；we must take synthetic measurements to accelerate the sustainable utilization of land resource, including administrative, economical ,technological and ecological policies.

青藏高原东缘几种树苗对增强紫外线-B和氮供应的响应

大气臭氧的损耗导致了地球表面具有生物学效应的紫外线-B(UV-B)辐射增强。同时，大气成分变化中除了UV-B辐射增强外，氮沉降是一个新近出现而又令人担忧的环境问题，其来源和分布正在迅速扩展到全球范围，并不断向陆地和水生生态系统沉降。本试验在四川省境内的中国科学院茂县生态站内进行，以云山、冷杉、色木槭和红椋子幼苗为模式植物，从生长形态、光合作用、抗氧化能力和矿质营养等方面研究了青藏高原东缘4种树苗对全球变化－增强UV-B辐射和氮供应(氮沉降)的响应。该试验为室外盆栽试验，包括四个处理：(1)大气UV-B辐射+无额外的氮供应(C)；(2)大气UV-B辐射+额外的氮供应(N)；(3)增强UV-B辐射+无额外的氮供应(UV-B)；(4)增强UV-B辐射+额外的氮供应(UV-B+N)。其目的：一方面有助于丰富我国对全球变化及区域响应研究的全面认识，进一步完善在全球气候变化条件下臭氧层削减和氮沉降对陆地生态系统影响的内容；另一方面，在一定程度上有助于我们更好的理解在全球变化下森林更新的早期过程。具体结果如下： 增强的UV-B辐射在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有显著的影响。UV-B辐射增强对幼苗的影响不仅与物种有关，而且，还与氮营养水平相关。总体表现为，高的UV-B辐射导致了色木槭和红椋子幼苗叶片的皱缩和卷曲，并降低了色木槭幼苗的叶片数和叶重，在额外的氮供应下，云杉、冷杉和红椋子的叶重也显著地降低了；色木槭和红椋子幼苗叶片的解剖结构受到了增强的UV-B辐射的影响，增强的UV-B辐射显著地降低了色木槭叶片的栅栏组织厚度，提高了红椋子叶片的厚度；增强的UV-B辐射显著地降低了4种幼苗的单株总生物量、植物地下部分的生长、总叶绿素含量 [Chl (a + b)]、净光合速率和最大量子产量(Fv/Fm)，提高了4种幼苗叶片的膜脂过氧化(MDA含量)，改变了植物体不同器官中的矿质元素含量；增强的UV-B辐射提高了冷杉、色木槭和红椋子叶片中的过氧化氢含量(H2O2)、超氧负离子(O2-)生成速率，在额外的氮供应下，云杉叶片中的活性氧含量也显著地提高了；在无额外的氮供应条件下，增强的UV-B辐射显著地提高了4种幼苗叶片中的UV-B吸收物质、脯氨酸含量和抗氧化酶的活性(SOD、POD、CAT、GR和APX)。在额外的氮供应条件下，UV-B辐射的增强却显著地降低了冷杉叶片中脯氨酸含量和红椋子叶片中UV-B吸收物质含量，但是，在4种幼苗叶片中，5种抗氧化酶的活性对UV-B辐射的增强没有明显的规律性，增强的UV-B辐射显著地提高了云杉叶片中的POD、SOD和GR的活性，提高了冷杉叶片中的POD和GR活性，提高了色木槭叶片中的POD、SOD和CAT活性和红椋子幼苗叶片中的POD和SOD活性。从这些结果可知，植物在遭受高的UV-B辐射导致的过氧化胁迫时，植物体内形成了一定的保护机制，但是，这种保护不能抵抗高的UV-B辐射对植物的伤害。 额外的氮供应在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有一定的影响，不同幼苗对额外的氮供应响应不同，并且受到UV-B辐射水平的影响。在当地现有的UV-B辐射水平下，额外的氮供应显著地提高了幼苗的单株总生物量、植物地下部分的生长、Chl (a + b)、净光合速率(红椋子除外)、UV-B吸收物质(冷杉除外)、脯氨酸含量(红椋子除外)和部分抗氧化酶的活性，降低了H2O2的含量、O2-的生成速率和MDA含量(红椋子除外)，改变了植物体内部分矿质元素含量，显著地提高了云杉和冷杉叶片中的Fv/Fm。这些指标总体表明，在当地现有大气UV-B辐射水平下，额外的氮供应对植物的生长和发育是有利的。在增强的UV-B辐射水平下，4种幼苗的生长形态和光合大部分指标都没有受到额外氮供应的影响，额外的氮供应提高了红椋子幼苗的单株总生物量和Chl (a + b)含量，提高了冷杉和色木槭叶片中的活性氧含量和MDA含量，却降低了红椋子叶片中的活性氧含量；额外的氮供应也提高了云杉、色木槭和红椋子叶片中UV-B吸收物质和脯氨酸含量，降低了冷杉叶片中UV-B吸收物质和脯氨酸含量；在抗氧化酶活性方面，额外的氮供应降低了云杉、冷杉叶片中5种抗氧化酶的活性和红椋子叶片中POD和GR的活性，提高了色木槭叶片中的POD和SOD的活性；4种幼苗植物体内的矿质元素含量对额外的氮供应没有显著的规律性。从这些结果可知，在高的UV-B辐射下，额外的氮供应提高了云杉、冷杉和色木槭幼苗对高的UV-B辐射的敏感性，然而，额外的氮供应却促进了红椋子幼苗的生长，原因可能是，在高的UV-B辐射下，额外的氮供应增加了红椋子叶片的厚度、叶重和叶片数，降低了叶片中活性氧含量的结果。表明在高的UV-B辐射水平下，额外的氮供应降低了红椋子幼苗对高的UV-B辐射的敏感性。 在全球变化的趋势下，UV-B辐射增强和氮沉降可能同时存在，我们的研究表明，与大气UV-B辐射+无额外的氮供应处理相比，增强UV-B辐射+额外的氮供应处理显著地降低了幼苗的单株总生物量(红椋子除外)、Chl (a + b)、净光合速率、Fv/Fm(冷杉除外)和MDA含量(红椋子除外)，提高了活性氧含量 (云杉除外)、UV-B紫外吸收物质含量(冷杉除外)、脯氨酸含量和部分抗氧化酶的活性，改变了植物体不同器官中的矿质元素含量。结果表明，在当地现有条件下，全球变化(UV-B辐射增强和氮沉降)对云杉、冷杉和色木槭幼苗的生长是不利，尽管植物体内一些抗氧化性指标提高了，然而，却对红椋子幼苗的单株总生物量的累积没有显著的影响。 The depletion of the ozone led to the increase of ultraviolet-B (UV-B) with biological effects in the earth’s surface. At the same time, except for enhanced UV-B radiation, nitrogen deposition was an anxious environmental problem at present, rapidly expanding to the global scope and continuously depositing to land and aquatic ecosystem. The experiment was conducted in Maoxian Ecological Station of Chinese Academy of Sciences, Sichuan province, China. Picea asperata, Abies faxoniana, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings were selected as model plants to assess the effects of enhanced UV-B radiation and supplemental nitrogen supply on growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings in east Qinghai-Tibetan Plateau. The experiment was potted outdoor, including 4 treatments: (1) ambient UV-B without supplemental nitrogen (control, C); (2) ambient UV-B with supplemental nitrogen (N); (3) enhanced UV-B without supplemental nitrogen (UV-B); (4) enhanced UV-B with supplemental nitrogen (UV-B+N). One hand, it was helpful for enriching our country to comprehensive understanding of the researches in the global change and the region response, further perfecting the effects of the depleted ozone layer and nitrogen deposition on land ecosystem under the global change; the other hand, it was favorable for us to better understanding of the early process of forest renews under the global change. The results were as follows: Enhanced UV-B radiation had significant effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings. The effects of enhanced UV-B on plants were not only related with species, but also related with nitrogen nutrient level. Generally, the increase of UV-B radiation led to the shrinkage and curl of leaves in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, and reduced the number of leaf and leaf weight of Acer mono Maxim seedlings, under supplemental nitrogen supply, leaf weight of Picea asperata, Abies faxoniana and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings significantly also reduced; the anatomical features of leaf in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings affected by enhanced UV-B radiation, the increase of UV-B radiation markedly reduced the palisade tissue thickness of Acer mono Maxim leaf and enhanced the leaf thickness of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings; the enhanced UV-B radiation significantly reduced total biomass per plant of 4 species seedlings, the growth of the underground parts, Chl (a + b), net photosynthetic rate and maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA content) and changed the content of mineral elements in different parts of plants; the enhanced UV-B radiation also increased the rate of superoxide radical (O2-) production and hydrogen peroxide (H2O2) content in leaves of Abies faxoniana, Acer mono Maxim, Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, under supplemental nitrogen supply, the reactive oxygen species in leaves of Picea asperata seedlings also significantly increased by enhanced UV-B radiation; under without supplemental nitrogen supply, enhanced UV-B radiation evidently induced an increase in UV-B absorbing compounds, proline content and the activities of antioxidant enzymes (SOD, POD, CAT, GR and APX) of leaves in 4 species seedlings. Under supplemental nitrogen supply, enhanced UV-B radiation induced a decrease in proline content of leaves in Abies faxoniana seedlings and UV-B absorbing compounds of leaves in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, but, there were no obvious rules in the activities of five antioxidant enzymes of 4 species seedling leaves to enhanced UV-B radiation, enhanced UV-B radiation significantly increased the activities of POD, SOD and GR in Picea asperata leaves, the activities of POD and GR in Abies faxoniana leaves and the activities of POD, SOD and CAT in Acer mono Maxim leaves. The results indicated that some protective mechanism was formed when plants were exposed to enhanced UV-B radiation, but the protection could not counteract the harm of high UV-B radiation on plants. Supplemental nitrogen supply had some effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits. The response of 4 species seedlings was different to supplemental nitrogen supply, and was affected by UV-B levels. Under local ambient UV-B radiation, supplemental nitrogen supply significantly increased the total biomass per plant, the growth of underground parts, Chl (a + b), net photosynthetic rate (except for Acer mono Maxim seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and the activities of some antioxidant enzymes, and reduced H2O2 content, the rate of O2- production and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and changed the content of mineral elemental in different parts; supplemental nitrogen supply also evidently increased Fv/Fm in Picea asperata and Abies faxoniana seedlings. These results indicated that supplemental nitrogen supply was favorable for the growth of plants under local ambient UV-B radiation. Under enhanced UV-B radiation, mostly parameters in growth and morphology of 4 species seedlings were not affected by supplemental nitrogen supply. Supplemental nitrogen supply increased the total biomass per plant and Chl (a + b) of Swida hemsleyi (Schneid. et Wanger.) Sojak seedling, increased the reactive oxygen species and MDA content in Abies faxoniana and Acer mono Maxim leaves, and reduced the reactive oxygen species in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves; supplemental nitrogen supply also increased UV-B absorbing compounds and proline content in Picea asperata, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, decreased UV-B absorbing compounds and proline content in Abies faxoniana leaves; in the activities of antioxidant enzymes, supplemental nitrogen supply significantly reduced the activities of antioxidant enzymes in Picea asperata and Abies faxoniana leaves and the activities of POD and GR in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, and increased the activities of POD and SOD in Acer mono Maxim leaves; the content of mineral elements in 4 species seedlings was no significantly rule to supplemental nitrogen supply. We knew from the results, under enhanced UV-B radiation, supplemental nitrogen supply made Picea asperata, Acer faxoniana and Acer mono Maxim seedlings more sensitivity to enhanced UV-B radiation, however, accelerated the growth of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings. The reason was probably that supplemental nitrogen supply increased the leaf thickness, leaf weight and leaf number, reduced the reactive oxygen content of leaf in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings grown under high UV-B radiation. This showed that supplemental nitrogen supply reduced the sensitivity of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings to high UV-B radiation. Under the tendency of the global change, enhanced UV-B radiation and nitrogen deposition may probably coexist. The results showed, compared with the treatment of ambient UV-B radiation without supplemental nitrogen supply, the treatment of enhanced UV-B radiation with supplemental nitrogen supply significantly reduced the total biomass per plants (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), Chl (a + b), net photosynthetic rate, Fv/Fm and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), and increased reactive oxygen content (except for Picea asperata seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content and part antioxidant enzymes, and changed the content of mineral elements of different parts. The results indicated that the global change (enhanced UV-B and nitrogen deposition) were not favorable for the growth of plants under local ambient UV-B radiation and nitrogen nutrient level,, though increased some antioxidant indexes, however, the treatment of enhanced UV-B with supplement nitrogen supply did not significantly affect on the biomass accumulation of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings.

CO2浓度升高对红桦幼苗生理与生长的影响

大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库，约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦（Betula albosinensis Burk.）是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物，采用人工模拟的方法，研究CO2浓度升高对不同种内竞争强度（种群水平）下红桦幼苗的生理特征、生长、干物质积累及其分配的影响，探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征，为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下： 1）在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后，叶片出现“光合适应”现象。与对照相比，低种植密度（28株m-2）和高种植密度（84株m-2）条件下的红桦幼苗净光合速率（A）、气孔导度（gs）、蒸腾速率（E）、表观量子产量（AQY）和羧化速率（CE）显著降低，而水分利用效率（WUE）则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物（TNC）浓度，结果是红桦幼苗的比叶面积（SLA，cm2 g-1）显著降低。 2）与对照相比，CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率（R GR）显著提高，尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积，结果是单株红桦幼苗的总叶面积比对照显著增加。 3）CO2浓度升高处理显著增加红桦幼苗干物质积累（尤其是细根生物量），改变了红桦幼苗生物量的分配格局。与对照相比，CO2浓度升高处理的红桦幼苗叶重比（LWR）、叶面积比（LAR）、叶根重比（Wl/Wr）和源汇重比（leaf weight to non-leaf weight ratio, Wsource/Wsink）显著下降（高种植密度的LWR除外），而根冠比（R/S）则显著增加。在两种种植密度条件下，CO2浓度升高显著增加红桦幼苗根生物量的分配比率，显著降低叶片的生物量分配比率，对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之，长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降，并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照，而后期迅速下降，说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比，表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration（[CO2]）has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.

环境CO2浓度和群落密度对红桦(Betula albosinensis Burk.)幼苗碳氮吸收和分配的影响

当前大气CO2浓度升高是全球变化的主要趋势之一，CO2浓度升高还会引起全球变暖等其它环境问题，因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种，在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验，研究了CO2浓度倍增环境下，不同密度水平红桦碳氮固定、分配可能发生的改变，并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应，为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下： 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长，同时也改变生物量在体内的分配格局，主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异，它们对CO2浓度升高的反应程度不同，但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应，也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度，显著增加红桦幼苗的群体生物量，从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度，对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加，但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理，这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明，红桦在升高CO2浓度的条件下，会作出积极的响应，从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高，植株各器官N、P含量降低，但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高，N、P向主茎和根的分配增加，向叶片的分配减少，主要是由于前者在总生物量中的比重增加，而后者减少了。(3) CO2浓度升高，氮磷利用效率(NUE和PUE)提高，氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后，亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加，后期积累速率下降。(3) 升高CO2浓度以后，土壤呼吸显著增强；土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的；所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期，随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势，同时MDA（丙二醛）含量下降，保护酶SOD（超氧化物岐化酶）、CAT（过氧化氢酶）活性升高。由此说明，升高CO2浓度有利于减缓生长季节后期叶片的衰老，使叶片维持较高的光合速率，也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.