19 resultados para SLA-Technologie
em Chinese Academy of Sciences Institutional Repositories Grid Portal
Resumo:
:传统的强调抗原多态性的SLA 分型方法主要是血清学、细胞学和生物化学的方法,随着分子生物学技术发 展,各具特点的SLA 分型技术不断涌现,如PCR2RFL P 法、PCR2SSCP 法、微卫星(MS) 法、DNA 序列的测定等,基于 强调SLA 的抗原多态性的分型和强调抗原保守性的功能学上的新分型技术(如SLA 超型和超基序) 进行了详细探 讨,比较了各类方法的优缺点,指明了未来SLA 分型的发展趋势。此外还指出了现行参考教材血清学SLA 单倍型 的编写错误以及重点强调了SL A2DQB 基因的准确分型技术必须借鉴于克隆测序。
Resumo:
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。
Resumo:
油蒿(Artemisia ordosica Krasch.)是内蒙古鄂尔多斯高原特有的半灌木,构成该地区沙地优势植被类型。主要分布在固定、半固定沙丘,同时在流动沙丘也有少量分布。它在当地经济价值、防风固沙环保方面均处于无以取代的地位。在毛乌素沙地沙漠化日益扩大的严峻态势下,研究其群落地上、地下过程对生境变化的响应不仅对维持干旱、半干旱区生态系统稳定的管理措施上有所帮助,而且也有助于了解全球变化背景下物种对环境条件的长期变化适应策略。 为此,本项研究以毛乌素沙地为研究区域,利用异速生长关系确立不同生境油蒿生物量最佳回归方程,并调查、比较了毛乌素沙地固定沙丘、半固定沙丘和流动沙丘三个生境油蒿灌丛地的生物量、土壤和植被的碳储量、生产力和细根周转、土壤微生物生物量碳、氮和土壤呼吸。具体结果如下: 1. 建立并比较了油蒿枝、株两个水平上各部分(不含细根)生物量异速生长关系式,其中枝形态指标(枝直径BD、枝长BL、叶枝长LBL)与油蒿叶、枝、果各部分生物量的异速关系最好;株水平上冠层面积CA与其叶、枝干、果、粗根各部分生物量的回归效果较好。不同生境生物量与其生长变量的异速生长关系存在差异。2004年调查的油蒿灌丛生物量从固定沙丘、半固定沙丘到流动沙丘分别是354.8,178.3和30.4 g m-2;各部分(叶、枝干、果、粗根、不同径级细根的)生物量都呈递减趋势。地下根与总生物量比值排序为固定>半固定>流动沙丘。不同生境细根生物量垂直分布存在差异,在固定沙丘根可至100 cm,半固定沙丘达90 cm,而在流动沙丘仅为60 cm,这些结果有助于使了解不同生境中的相同物种如何通过自身形态及其生物量调整来适应生境的差异。 2. 不同生境油蒿灌丛地植被碳储量和土壤碳储量在P < 0.05水平上差异显著,其中固定沙丘植被碳储量和土壤碳储量分别为224.04和7521 g C m-2,半固定沙丘是119.27和3029 g C m-2,流动沙丘是16.83和2300 g C m-2。可见沙区土壤有机碳远大于植被碳量。 3. 利用最大值减最小值方法、标准取样法和内生长土芯法研究了不同生境油蒿灌丛地的地上、粗根生产力和地下细根生产力。发现各生境生产力、细根周转都随着植被盖度增加而增加,地下根生产力与总生产力之比随着植被盖度增加而减少;不同生境油蒿灌丛地生产力在P < 0.05水平上存在显著差异,2005年总生产力范围在18.23-293.82 g m-2 yr-1之间;细根总周转率在0.16-0.54 yr-1之间。 4. 利用异速生产关系确立不同生境不同水平上油蒿叶面积的最佳回归关系式并对不同生境的比叶面积(SLA)进行了比较,其中枝水平上各生境叶面积与枝直径、枝叶长、枝长相关关系在P < 0.001水平上显著;株水平上各生境叶面积与株高、冠层面积相关关系在P < 0.001水平显著;从固定沙丘、半固定沙丘到流动沙丘SLA由大变小,这可能与生境养分差异有关。 5. 不同生境油蒿灌丛地土壤微生物碳、氮和土壤呼吸范围分别在117.99-153.99 mg kg-1、1.49-3.31mg kg-1和0.54-1.96 μmol m-2 s-1之间,它们从固定沙丘、半固定沙丘到流动沙丘依次下降。
Resumo:
本文选取不同放牧率下的优势植物为研究对象,主要从植物解剖结构和化学成分方面,对内蒙古典型草原和北美混合普列里草原的放牧演替机制进行探讨,这将有助于进一步揭示放牧演替过程及其主要植物对放牧的适应机理。具体研究结果如下: 1. 长期不同放牧率的放牧(内蒙古典型草原区放牧15年和混合普列里草原放牧19年)对所有物种(IMGERS的羊草(Leymus chinensis)、冰草(Agropyron cristatum)、糙隐子草(Cleistogenes squarrosa)、扁蓿豆(Melissitus rutenica)、小叶锦鸡儿(Caragana microphylla)、冷蒿(Artemisia frigida)和星毛委陵菜(Potentilla acaulis)和CGREC的Artemisia frgida ,Poa pratensis, Agropyron smithii,Solidago rigida, Helianthus rigidus和Symphoricarpos occidentalis)叶片的角质层厚度、表皮细胞面积、叶肉细胞面积、栅栏/海绵组织厚度、叶片厚度、中脉厚度均产生显著影响;放牧显著影响了两个研究区不同生活型功能群植物叶片的角质层厚度、表皮细胞面积、叶肉面积、栅栏/海绵组织和中脉厚度。内蒙古典型草原研究区草本植物功能群的叶片下角质层厚度、栅栏/海绵组织厚度显著大于灌木功能群,而美国混合普列里草原研究区,草本植物功能群的叶片下角质层厚度、表皮细胞面积、叶肉细胞面积、栅栏/海绵组织厚度、叶片厚度、中脉厚度均显著大于灌木功能群。 2. 内蒙古典型草原研究区放牧率显著影响了糙隐子草和小叶锦鸡儿的比叶面积(SLA);在美国混合普列里草原研究区,放牧率显著影响了冷蒿SLA。但对生活型功能群的SLA影响不显著。 3. 内蒙古典型草原研究区放牧对物种叶片叶绿素含量、纤维素含量影响显著,放牧仅显著增加了扁蓿豆叶片的含氮量。美国混合普列里草原研究区物种叶片的全碳、干物质、酸性洗涤纤维、叶绿素a+b含量受放牧率的显著影响。两研究区放牧率对叶片叶绿素a+b含量影响显著。放牧率也显著影响了不同生活型功能群的全碳含量和叶绿素a+b含量。 4. 两研究区叶片表皮细胞面积和叶肉细胞面积无牧和重牧下显著正相关,表皮细胞面积和叶片厚度在轻牧下显著正相关,叶肉细胞面积和叶片厚度在无牧、中牧和重牧下显著正相关,比叶面积和叶绿素a+b含量在轻牧下显著正相关。扁蓿豆叶片的表皮细胞面积和叶肉细胞面积间存在显著的正相关,其全碳含量和叶绿素a+b含量间存在显著的正相关;Artemisia frgida叶片的角质层厚度和叶片厚度间显著正相关,Poa pratensis叶片的角质层厚度和表皮细胞面积间显著负相关,Solidago rigida叶片表皮细胞面积和光合速率显著正相关。Helianthus rigidus叶片叶肉细胞面积和叶片厚度显著负相关。
Resumo:
养分回收是植物衰老时,养分从衰老组织向活的组织体转移的一种过程。该过程延长了养分在植物体内的滞留时间,提高了植物保持养分的能力,因此是植物适应养分贫瘠生境的策略之一。全球气候变化,包括降水格局改变和氮(N)沉降增加,改变了生态系统正常的生物地球化学循环,因此可能会对植物的养分回收特性产生影响。研究不同N、磷(P)、水梯度上,植物物种养分回收特性的响应格局,对于预测N沉降增加和降水格局改变对物种养分利用策略的潜在影响,具有一定的理论与实践意义。本研究以中国科学院植物研究所多伦恢复生态学实验站长期施N肥实验(0,1,2,4,8,16,32,64 g N m-2 yr-1等8个水平)为研究平台研究了克氏针茅(Stipa krylovii)群落中优势植物养分回收随N素添加梯度的变化,同时结合三个盆栽控制实验(施N肥实验:0,0.5,1,2,4,8,16,32,64,128 g N m-2等10个水平;施P肥实验:梯度同施N肥实验;控水实验:3600,4000,4500,5143,6000,7200,9000,12000,18000,36000 mL pot-1等10个水平),主要探讨了羊草(Leymus chinensis)养分回收效率(从衰老组织中回收转移的养分百分数,RE)和养分回收度(以枯叶中养分浓度衡量,RP)以及其它叶片养分特性(绿叶养分浓度和比叶面积SLA)对环境因子改变的响应格局。同时,我们还调查了枯叶C:N比和C:P比等参数,研究环境因子改变对凋落物分解的影响。 1)连续4年施N肥显著降低了五个物种叶片N素回收度(NRP),对P素回收度(PRP)的影响在各物种间差异较大,但低N提高了多数物种的PRP;物种间,冷蒿(Artemisia frigida)RP(枯叶N和P浓度分别为14.3±2.0 mg g-1和0.68±0.09 mg g-1)最低,砂韭(Allium bidentatum)(N:5.2±0.2 mg g-1,P:0.12±0.01 mg g-1)最高。沿施N梯度,N素回收效率(NRE)的变化趋势在物种间差异较小但在方法间(叶干重水平,叶面积水平和单株水平)差异较大,而P素回收效率(PRE)的变化在物种间和方法间差别都较大。叶干重水平和叶面积水平上,NRE在四个物种中表现出显著降低的趋势,PRE只在糙隐子草(Cleistogenes squarrosa)和星毛委陵菜(Potentilla acaulis)中显著降低,其它三个物种变化不显著。单株水平上,所有物种NRE(除了克氏针茅)和PRE均与施N量梯度无显著性关系。物种间,砂韭的RE最高(>80.0%),冷蒿和星毛委陵菜最低(<60.0%)。方法间,叶片水平上的RE均高于单株水平上。沿施N肥梯度,两个禾本科物种SLA无显著变化规律,而其它三个物种SLA表现出先增加后变化不大的趋势。物种间,最高和最低的SLA分别表现在冷蒿和克氏针茅。沿施N肥梯度,五个物种C:N比呈先显著降低后缓慢降低的趋势。物种间,最大和最小的C:N比分别出现在砂韭和冷蒿。 2)盆栽施N肥实验中,一定范围内,施N肥显著提高了羊草地上地下生物量、SLA和绿叶N浓度,显著降低了C:N比、NRP、NRE和PRE,但对绿叶P浓度、叶片PRP和C:P比无明显影响。平均枯叶N浓度和枯叶P浓度分别为16.2 mg g-1和1.01 mg g-1,平均NRE和PRE分别为46.1﹪和58.1﹪。10月时,地下生物量和养分积累均高于地上部分。 3)盆栽施P肥实验中,一定范围内,施P肥显著地提高了羊草地上地下生物量、SLA、绿叶N浓度和绿叶P浓度,显著降低了C:N比、C:P比、NRP、PRP和PRE,但对NRE无显著影响。平均枯叶N浓度和枯叶P浓度分别为9.9 mg g-1和7.43 mg g-1,平均NRE和PRE分别为58.2﹪,平均PRE为56.1﹪。10月时地下部分生物量和N库积累均高于地上部分,而P库在两个部分间差别不大。 4)盆栽控水实验中,一定范围内,供水量增加显著增加了羊草地上地下生物量、SLA、NRP、PRP、PRE、C:N比和C:P比,显著降低了绿叶N浓度,但对绿叶P浓度和NRE无显著性影响。平均枯叶N浓度和枯叶P浓度分别为10.4 mg g-1和0.32 mg g-1,平均NRE和PRE分别为54.4﹪和76.8﹪。10月时,地下部分生物量和养分积累均高于地上部分。 以上结果表明,N、P和水分因子的改变影响了植物生物量和养分分配、叶片养分特性、养分回收能力以及枯叶分解质量等,且不同梯度影响程度也不同。因此,未来全球变化包括N沉降增加和降水格局改变可能影响植物养分利用策略和凋落物分解特性,进而可能对植被-土壤系统养分循环产生影响。
Resumo:
植物性状是植物与环境长期互作过程中表现出的内在或外部的特征。目前植物生态学界关于植物性状的研究,主要包括以下三个方面的研究内容:植物性状与环境因子间的关系,反映植物性状对环境变化的响应或适应机制;植物性状与生态系统功能的关系,以更好地揭示生态系统功能的内在驱动机制;植物性状间的相关关系和消长对策,反映植物性状的协同进化机制。 本试验所在地区内蒙古多伦十三里滩为典型克氏针茅草原,水分为当地生态系统的主要限制因素。我们在当地的围封样地内,采用人工模拟生长季降雨变化梯度的方法,研究了内蒙古典型草原几种常见植物叶片功能性状对模拟降雨量梯度的响应、性状之间的协变关系以及植物性状与地上部分生物量间的关系。研究工作在2005-2006年期间进行。观测的叶片性状包括叶片干物质含量(LDMC),比叶面积(SLA)和叶片氮含量(LNCmass, LNCarea),以地上部分生物量作为年净初级生产力(ANPP)估计,降雨量梯度模拟了两个水平,即对照(接收正常降雨)和增雨。主要结论如下:1) 叶片性状在降雨量梯度上的变化趋势在物种水平上表现出多样化的特点,性状的变化趋势是否有显著的格局取决于物种或特定性状。2) 叶片性状对水分梯度的响应在功能群水平上表现出各自的特点,Grass功能群对水分梯度的响应敏感性较低,Forb居中,Legume表现出较强的敏感性,表明Grass功能群对水分的耐受性比较强,这在一定程度上解释了当地克氏针茅草原上目前以禾草类为主的现状,对物种之间的替代有一定的指示性。而群落水平上探讨性状在环境梯度上的变异趋势,研究结论存在较大的不确定性,主要是难以区分环境和系统发育背景对植物性状的影响。3)叶片功能性状间的协变关系,不同物种因其自身发生背景不同,在不同环境背景下总体表现出一致的变化规律,但也存在例外。相对而言,对功能性状间协变关系的研究在物种水平或功能群水平上更有意义,而在群落水平上所得结论,可能会掩盖了群落内部不同物种或不同功能群间的信息。4)叶片功能性状与草原地上部生物量的关系在功能群水平上呈显著正相关关系,再次验证了土壤含水量的变化同土壤氮元素的动态,共同影响了植物的氮利用策略,进一步影响了初级生产力等和氮元素直接相关植物性状参数。在群落水平上叶片功能性状与生物量的之间并没有表现出可遵循的规律性,说明叶片功能性状对生物量的作用更多地局限在群落内部的组织层次上发挥作用。5)本研究的虽然从不同的组织层次上探讨了叶片功能性状对环境、性状之间的协变关系以及与地上部分生物量间的关系,由于植物性状变化幅度可能与环境梯度密切相关,如果模拟环境梯度不够大,可能会影响结论的普遍性。此外,本次野外控制试验时间较短,数据量偏少,本文的结论尚需要进一步验证。
Resumo:
CO2浓度升高和气候变暖已成为不可逆转的现实,而具有强氧化性的臭氧浓度升高也被认为是全球变化的重要组成部分,已经或将对植被生长产生严重威胁。冬小麦(Triticum aestivum L.)生长发育期内受到环境变化的影响相对较大。小麦是对臭氧最为敏感的作物之一,华北地区冬小麦灌浆期容易发生臭氧污染。本研究以冬小麦为试材,研究了CO2浓度加倍和2oC增温对中国北方不同年代推出的冬小麦品种幼苗生长的效应;研究了不同小麦品种对花后短期臭氧胁迫的反应;分别研究了干旱、增温与臭氧污染对冬小麦旗叶光合和产量的交互影响。研究结果如下: 1) CO2浓度加倍和温度升高加快冬小麦幼苗的生长速率。CO2浓度加倍主要是促进了幼苗的分蘖(+1.54分蘖),而增温加快了发育进程(主茎叶片数比对照多1叶)。不同品种对CO2浓度加倍和增温的响应存在着差异,但与育种年代没有显著相关关系。从20个品种的总体情况来看,CO2浓度加倍和2.0℃增温对冬小麦幼苗的生长具有叠加效应。生长较慢的冬小麦品种对CO2浓度加倍的反应越大。 2) CO2浓度加倍显著提高了小麦新展开叶的净光合速率(Psat),显著降低了气孔导度(gs)和蒸腾速率(E),从而显著提高了瞬时水分利用效率(ITE),但不同小麦品种对CO2浓度加倍的反应与其育种年代没有明显的相关性;增温对冬小麦叶片Psat的效应不显著,但显著降低了gs,而E显著增大,从而导致ITE显著降低,其中,Psat对增温的反应与品种育种年代呈显著正相关(R=0.525, p<0.05),并与叶片比叶面积(SLA)显著正相关(R=0.45, p<0.001)。CO2浓度加倍和增温同时处理时对Psat、gs和ITE的效应显著大于CO2浓度加倍处理。 3) 冬小麦灌浆初期,短时间的臭氧胁迫对旗叶的同化能力带来了显著的负效应,而且光合性能最大的叶片中部受到的抑制最为严重。同化能力的下降导致最终产量的降低。然而,不同品种对臭氧的敏感性存在着很大差异。臭氧对具有较高收获指数和产量的当代品种-烟农19的效应(-19%,p<0.01)明显大于产量较低的老品种-农大311(-8%,p<0.05),主要表现在叶片可见伤害程度较大,旗叶净光合速率受到的负效应较大,籽粒重量以及穗粒数均显著降低。 4) 短期高浓度臭氧胁迫对处于中度干旱胁迫下的小麦植株产生了可见伤害(<20%),但伤害程度明显低于处于土壤含水量较高的植株(>30%)。短期臭氧处理显著降低了处于较高土壤含水量下的小麦旗叶Psat(-36%),干旱胁迫也显著降低了旗叶Psat(-34%),但臭氧仅使处于中度干旱胁迫下的植株旗叶Psat进一步降低了7.8%。gs的变化趋势与Psat的变化基本一致。臭氧处理结束并复水后,干旱处理植株的Psat与对照基本相同,而臭氧处理过的植株均明显低于对照,其中臭氧处理期间处于良好土壤含水量条件下的植株旗叶Psat显著低于对照,并且随着植株的衰老,其下降的速度明显快于其它处理,表明衰老速度加快。而中度干旱胁迫可减轻臭氧对小麦产量的危害程度。 5) 正常供水条件下,短期臭氧处理结束2天后,小麦旗叶Psat与对照植株的值接近,然而随着时间的延长,Psat下降的速度明显快于对照。臭氧处理结束后,植株若遭受干旱胁迫,旗叶Psat显著低于对照。虽然gs的变化趋势与Psat基本一致,但臭氧处理后光合的降低主要是由非气孔限制因素引起的。旗叶光合能力下降、叶片提前衰老,是臭氧处理导致产量显著降低的主要原因。穗粒数没有显著变化,而千粒重却显著降低。而臭氧处理后若遇干旱胁迫,旗叶的光合能力以及光合有效期进一步减少,导致不育小花数增多,因此籽粒产量进一步减少。 6) 灌浆期臭氧浓度升高显著降低了小麦叶片的光合能力,导致产量显著降低;2oC升温加快了植株的衰老,然而并没有对产量性状产生显著效应;温度和臭氧两因子对旗叶的气体交换参数具有交互效应,但对各产量性状均没有交互效应发生。 综合以上研究结果,不同品种对环境变化的响应存在着显著差异,意味着可以通过育种途径来减轻未来环境变化对农业生产的负面影响;灌浆期臭氧胁迫显著降低冬小麦的产量,环境因子可影响臭氧胁迫效应,但之间的交互作用比较复杂,有待进一步深入研究,以保证未来全球变化环境下的粮食安全。
Resumo:
随着Internet的飞速发展,符合J2EE规范的Web容器已经成为部署企业Web应用的主流平台.同时,企业Web应用的多样化和复杂性的增长,迫切地要求Web容器提供差分服务的支持.然而,传统的Web容器采用尽力而为的服务模型,无法提供差分服务支持.已有的研究采用准入控制、优先级调度等方法来为Web容器提供差分服务支持,但是它们只能提供单调、静态的差分服务策略.提出了一个基于自管理单元的Web容器DSWC,它能够根据SLA定义提供两级差分服务支持.此外,提出了一个自适应差分服务策略选择算法,它能够根据动态变化的运行环境自适应地选择差分服务策略.原型系统的实验结果显示,DSWC能够有效地为请求提供符合SLA定义的细粒度的差分服务支持.
Resumo:
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.
Resumo:
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.
Resumo:
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.
Resumo:
植物生长和生产力受到自然界各种形式的生物和非生物胁迫因子的影响。这些胁迫包括低温、高温、盐碱、干旱、洪水、重金属、虫害、病害和紫外线辐射等等。而人类活动大大加剧了这些胁迫所带来的影响。由于人类污染而导致臭氧层衰减以及由此产生的地球表面紫外辐射增强已经成为全球气候变化的一个主要方面。UV-B胁迫,甚至当前的辐射水平,所带来的影响已经引起科学工作者的广泛关注。 为了生存和繁殖,植物不得不面临环境中各种潜在胁迫所带来的负面影响。然而,植物生活型的不可移动性决定了其逃避胁迫的局限性。因此,绝大多数植物都是通过对胁迫作出反应,通过修复或者更新组织来降低伤害。而植物应对环境变化的能力则是由其生长模式的种属特异性和本身的遗传组成所决定。在自然界,植物常常同时面临多种胁迫,这些胁迫所引发的植物反应可能具有叠加、协同或者拮抗作用。沙棘是一种具刺、具有固氮功能的多年生雌雄异株灌木,广泛分布于亚欧大陆的温带地区和亚洲亚热带的高海拔地区。在中国,沙棘常常被用作植被恢复中的先锋树种而大量栽培。本文采用沙棘作为模式植物,试图探索木本植物对低温,UV-B辐射增强以及其与干旱的复合胁迫的响应以及沙棘对这些胁迫响应是否具有种群差异性。 对来自南北两个种群的沙棘进行短日照和低温处理,检测了其在抗寒锻炼和抗寒性发育过程中存在的性别差异。结果表明,短日照和低温都分别能够诱导抗寒锻炼的发生,而两者同时存在对所有实验植株抗寒性的大小具有叠加效应。然而,短日照和低温所诱导的抗寒性在两个种群中都具有性别差异性,雄性植株比雌雄植株对短日照和低温更为敏感。同时,南北种群间也存在差异性,北方种群的植物比南方种群的植物对短日照和低温敏感,从而在短日照下抗寒锻炼的发生时间更早,低温诱导的抗寒性更大。短日照和低温诱导植物增加抗寒性的同时伴随着脱落酸的变化。脱落酸的变化因处理,种群和性别的不同而不同。这些生理反应表明不同的沙棘种群,不同的植株性别对同一环境胁迫可能存在不同的生存策略。 比较了来自高低两个海拔的沙棘种群对于干旱和UV-B辐射增强以及两者复合胁迫条件下的生理生态反应。干旱使两个种群中植株总的生物量,总叶面积,比叶面积,叶片含碳量,含磷量,木质素含量和碳氮比显著降低,使根冠比,粗根细根比和叶片脱落酸含量显著增加。干旱而非UV-B使得δ13C 值显著增加。但是,比较而言,来自高海拔的种群对干旱反应更为强烈,而来自低海拔的种群对UV-B更敏感。在UV-B辐射增强的处理下,干旱所诱导的脱落酸的积累被显著抑制。而且我们检测到在一些指标上存在显著的干旱×UV-B交互作用,如两个种群中在总生物量上,低海拔种群中在总叶面积,粗根细根比上,高海拔种群中在比叶面积,δ13C值,木质素含量上都存在明显的交互作用。这些结果表明这两个种群对胁迫具有不同的适应性反应,来自高海拔的种群比来自低海拔的种群更能够抵御干旱和UV-B胁迫。 室外实验表明,UV-B 去除/增补对沙棘高低两个海拔种群的影响都不大。对生物量的积累,植株高度以及一些常见的胁迫反应生理指标比如丙二醛、ABA 和游离脯氨酸都没有显著影响。UV-B 的效应比UV-A 大,植物反应在无UV 和仅有UV-A 的处理间没有什么区别。然而,UV-B 去除的两个处理和UV-B 存在的两个处理间存在显著区别。UV-B 使得两个种群都显著降低了比叶面积(SLA),但却使长期用水效率增加。但UV-B对光合色素和光合系统II 的影响不大。总体看来,来自低海拔的种群对UV-B 更为敏感。 Plant is adversely affected by various abiotic and biotic stress factors. These stressors includelow temperature, heat, salt, drought, flooding, heavy metal toxicity, wounding by herbivores,infecting by pathogenic microorganisms, ultraviolet (UV) radiation and so on. Variousanthropogenic activities have accentuated the existing stress factors. One of the mostimportant aspects of global change is that of stratospheric ozone depletion caused by seriousanthropogenic pollution and the resulting increase in UV radiation reaching the surface of theEarth. Scientists have become concerned about the effects that considerable UV-B stress, evenat current levels. In order to survive and reproduce, plants have to be able to cope with lots of potentiallyharmful stress factors that are almost constantly present in their environment. Most plants’responses under stress are to neutralize the stress, repairing the damage or regrowing newtissue rather than to avoid it due to their sessile life style. The plant defense capacity dependson plant-specific modular growth patterns and genetic make-up that allows for flexibleresponses to changing environments. Plants usually encounter several stresses simultaneouslyunder field conditions, and the stresses may cause a variety of plant responses, which can beadditive, synergistic or antagonistic. Sea buckthorn (Hippophae rhamnoides L.), a thorny nitrogen fixing deciduously perennialshrub, which is widely distributed throughout the temperate zones of Asia and Europe and thesubtropical zones of Asia at high altitudes. It has been widely used in forest restoration as thepioneer species in China. In this paper, we used sea buckthorn as a model, tried to get some understand of how plants fight low temperature, enhanced UV-B radiation level and thatcombination of drought. And also, want to know whether does there exist some populationspecific responses to such stressors. Sexual differences in cold acclimation and freezing tolerance development of two contrastingsea buckthorn (Hippophae rhamnoides L.) ecotypes from northern and southern regions inChina were recorded after exposure to short day photoperiod (SD) and low temperature (LT).The results demonstrated that cold acclimation could be triggered by exposing the plants toSD or LT alone, and that a combination of both treatments had an additive effect on freezingtolerance in all plants tested. However, development of freezing tolerance was dependent onthe sex of plants under SD and LT, the males were clearly more responsive to SD and LT thanthe females in both ecotypes studied. On the other hand, development of freezing tolerancewas also ecotype-dependent, the northern ecotype was more responsive to SD and LT than thesouthern ecotype, resulting in earlier cold acclimation under SD and higher freezing toleranceunder LT. Moreover, development of freezing tolerance induced by SD and LT wasaccompanied by changes in ABA levels. These alterations in ABA levels were different indifferent treatments, ecotypes and sexes. Therefore, the differences in SD and LT-inducedphysiological responses showed that the different ecotypes and the different sexes mightemploy different survival strategies under environmental stress. Two contrasting populations from the low and high altitudinal regions were employed toinvestigate the effects of drought, UV-B and their combination on sea buckthorn. Droughtsignificantly decreased total biomass, total leaf area, specific leaf area,leaf carbon (C),phophous (P), lignin content and the ratio of C: N in both populations, and increasedroot/shoot ratio, fine root/coarse root ratio and abscisic acid content (ABA), in bothpopulations. Drought but not UV-B resulted in significantly greater carbon isotopecomposition (δ13C) values in both populations. However, the high altitudinal population wasmore responsive to drought than the low altitudinal population. The drought-inducedenhancement of ABA in the high altitudinal population was significantly suppressed in thecombination of drought and elevated UV-B. Moreover, significant drought × UV-B interactionwas detected on total biomass in both populations, total leaf area and fine root/coarse root inthe low altitudinal population, specific leaf area, δ13C value and leaf lignin content in the high altitudinal population. These results demonstrated that there were different adaptive responsesbetween two contrasting populations, the high altitudinal population exhibited highertolerance to drought and UV-B than the low altitudinal population. A field experiment was conducted to investigate effects of UV-B exclusion/supplementationon two altitudinal populations of sea buckthorn. UV-B exclusion or supplementation had littleeffects on both populations investigated. For instance, the total biomass, plant height andsome physiological index such as Malondialdehyde (MDA), ABA and free proline were notchanged significantly. The UV-B effects are more significant than that of UV-A, nodifferences were found between treatments of excluded UV and excluded UV-B. However,compared with treatments of UV-B exclusion (including absent of UV-B and all UV band),the present of UV-B (including near ambient environment and enhanced UV-B) significantdecreased specific leaf area, and increased long time water use efficiency as evaluated by δ13Cvalue. UV-B had little effects on photosynthetic pigments and Photosystem II (PSII). The lowaltitude population is more sensitive to UV-B than that of the high altitude population.
Resumo:
随着全球气候变暖和温室效应加剧,干旱和荒漠化成为威胁人类生存和发展的主要 灾害,许多被子植物对干旱胁迫的生理、生态和生化响应已逐步得以报道,但很少有开 展干旱胁迫对雌雄异株植物的影响方面的研究。由于这类植物在长期进化过程中已经在 生长、性比、生殖格局、空间分布、资源配置和生物量分配等方面形成了明显的性别差 异,因此,干旱胁迫必将对其雌雄植株产生不同的生理生态影响。本研究以青杨为模式 植物,采用植物生态、生理及生物化学等研究方法,系统研究青杨雌雄植株在常温、增 温以及喷施外源脱落酸的条件下对干旱胁迫的响应,揭示其在生长形态、生物量分配、 光合作用、用水效率和生理生化等方面的性别间差异。主要研究结果如下: 1. 青杨雌雄植株对干旱胁迫的综合响应。 与较好水分条件相比,干旱胁迫显著降低了青杨雌雄植株的光合作用和生长发育, 影响了许多生理生化过程,并导致雌雄植株在生长发育、气体交换、用水效率、膜脂抗 氧化和抗氧化系统酶活性方面表现出显著的性别间差异。在较好水分条件下,雌雄植株 之间在株高、基径、生物量、净光合速率、蒸腾速率、用水效率以及丙二醛、脱落酸和 游离脯氨酸等生化物质含量方面均无显著差异。但在干旱胁迫下,雄株在生长发育、气 体交换、水分利用效率、膜脂过氧化保护和抗氧化系统酶活性方面均显著高于雌株,表 现出比雌株更高的株高、基径、叶面积、总叶片数、总生物量、总色素含量、类胡萝卜 素含量、净光合速率、蒸腾速率、羧化效率、光系统II最大光化学效率、内在水分利用 效率、碳同位素组分、过氧化氢酶和过氧化物酶活性等,而在CO2补偿点、比叶面积、 叶绿素a/b、丙二醛、脱落酸和超氧化物歧化酶活性等指标上显著低于雌株。与雌株相比, 雄株表现出更高的干旱胁迫适应能力,而雌株的生长发育和生理生化过程更易遭受干旱 胁迫的影响。 2. 干旱胁迫下的青杨雌雄植株对增温处理的综合响应 与环境温度相比,增温在干旱胁迫前后均显著促进了雌雄植株的生长发育、气体交 换,降低水分利用效率,影响生化物质含量,并促使青杨雌雄植株之间在干旱胁迫下表 现出显著的差异。在较好水分条件下,增温导致雌株的株高、基径、叶面积、总叶片数、 总生物量和超氧化物歧化酶活性显著高于雄株,而用水效率、丙二醛、脱落酸和游离脯 氨酸、抗坏血酸过氧化物酶和过氧化物酶活性低于雄株。在干旱胁迫下,增温将导致雄 株的株高、基径、叶面积、总生物量、净光合速率、蒸腾速率、气孔导度、总色素含量、 相对含水量、过氧化氢酶和抗坏血酸过氧化物酶活性等显著高于雌株,而光系统II 最大 光化学效率、内在水分利用效率、碳同位素组分、丙二醛、脱落酸、游离脯氨酸和超氧 化物歧化酶活性显著低于雌株。与雄株相比,水分较好条件下的增温有利于促进雌株的 生长发育,并在生理生态特征上优于雄株。而干旱胁迫下的增温则加剧了水分胁迫强度, 致使雌株的生长发育遭受比雄株更多的负面影响。 3. 干旱胁迫下的青杨雌雄植株对喷施外源脱落酸处理的综合响应 与对照相比,在干旱胁迫下喷施外源脱落酸可显著增加青杨雌雄植株的生长发育、 气体交换、降低水分利用效率,影响了生化物质含量,并导致青杨雌雄植株之间在干旱 胁迫下表现出显著的生理生态差异。在干旱胁迫下,喷施外源脱落酸致使雌株的株高、 叶面积、叶干重、细根干重、总生物量、净光合速率、蒸腾速率、气孔导度、光系统II 最大光化学效率、非光化学淬灭系数、相对含水量、总光合色素、类胡萝卜素、脱落酸、 超氧化物歧化酶和过氧化物酶活性的增加量显著高于雄株,而根重比、根冠比、细根/ 总根、比叶面积、内在水分利用效率、碳同位素组分、丙二醛、脯氨酸、过氧化氢酶和 抗坏血酸过氧化物酶活性等指标的减少量上显著低于雄株。与对照相比,干旱胁迫下的 喷施外源脱落酸则一定程度能减缓植株遭受胁迫的压力,促进植株生长和气体交换,减 少了植株体内的过剩自由基数量,并促使雌株的生长发育和光合能力显著提高,增强其 抗干旱胁迫能力。 With development of global warming and greenhouse effect, drought and desertification have been became main natural disasteres in resent years. Studies on ecophysiological responses of most angiosperm species to environmental stress have been reported, but little is known about dioecious plant responses to drought stress. Since significant differences on growth, survival, reproductive patterns, spatial distribution, as well as resource allocation between males and females of dioecious plant have been formed during evolutionary process, sexual different ecophysiological responses should be caused by drought stress. In this experiment, Populus cathayana Rehd. was used as model plant to study the sex-related responses to drought by using the ecological, physiological and biochemical methods under normal atmospheric temperature, elevated temperatures and exogenous abscisic acid (ABA) application treatment respectively, and to expose the sexual differences in growth, biomass allocation, photosynthesis, water use efficiency and some biochemical material contents in the males and females of dioecious plant. The results are follows: 1. A large set of parallel responses of males and females of P. cathayana to drought stress Compared with well-watered treatment, drought significantly decreased growth and photosynthesis of P. cathayana individuals, affected some physiological and biochemical processes, and induced males and females to exhibit obvious sexual differences in growth, gas exchange, water use efficiency, lipid peroxidation protection and antioxidant defenses enzyme system. Under well-watered treatment, there were no significant sexual differences in height growth (HG), basal diameter (BD), dry matter accumulation (DMA), net photosynthesis rate (A), transpiration (E), water use efficiency (WUE), and malondialdehyde (MDA), abscisic acid (ABA) and praline (Pro). However, under drought stress, males were found to exhibit higher HG, BD, leaf area (LA), total leaf number (TLA), DMA, total chlorophyll contents (TC), carotenoids content (Caro), A, E, carboxylation efficiency (CE), the maximum efficiency of PSII (Fv/Fm), intrinsic water use efficiency (WUE ), carbon isotope composition (δ13C), catalase (CAT), peroxidase (POD) and lower CO2 compensation point (Γ), specific leaf area (SLA), chlorophyll a/b ratio (Chla/Chlb), MDA, ABA and superoxide dismutase (SOD) than females. The results suggest that males possess greater drought resistance than do females and females suffer more negative effect on growth and development, physiological and biochemical processes than males under drought stress. 2. A large set of parallel responses of drought-stressed males and females of P. cathayana to elevated temperatures Compared with environmental temperature, elevated temperature treatment significant increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents of P. cathayana individuals, and induced males and females to exhibit obvious differences under drought stress. Under good water condition, elevated temperature treatment caused females to show significant higher HG, BD, LA, TLN, DMA, SOD activity, and great lower WUE, MDA, ABA, Pro, ascorbate peroxidase (APX) and POD than do males. On contrary, under drought condition, elevated temperature treatment induced males to exhibit higher HG, BD, LA, DMA, A, E, stomatal conductance (gs), relative water content (RWC), CAT, APX activity but lower Fv/Fm, WUE, δ13C, MDA, ABA, Pro, SOD activity than do females. The results suggest that females will benefit from elevating temperature under good water condition by possessing better ecophysiological processes than that of males, but will suffer from greater negative effects than do males when grown under drought stress with elevated temperature treatment. 3. A large set of parallel responses of drought-stressed males and females of P. cathayana to exogenous ABA application Compared with controls, exogenous ABA application under drought greatly increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents in P. cathayana individuals, and induced males and females to exhibit obvious sexual differences under drought. Under drought stress, exogenous ABA application induced females to exhibit more increases in HG, LA, leaf weight (LW), fine root weight (FRW), DMA, A, E, g, Fv/Fm, non-photochemical quenching coefficient (qN), RWC, TC, Caro, ABA, SOD, POD s activity than males, but to show lower decreases in root/weight ratio (RWR), root mass/foliage area ratio (RF), fine root/total root ratio (FT), SLA, WUE, δ13C, MDA, Pro, CAT, APX than males. The results suggest that exogenous ABA application under drought stress will eliminate negative damages caused by drought stress at a certain extent,promote the growth and gas exchange of plant and decrease the number of superfluous 1O2 in plant cells of males and females of P. cathayana. Furthermore, exogenous ABA application promoted more drought resistance in females than in males by increasing more growth and photosynthetic capacity in females under drought stress.