36 resultados para Betula albosinensis
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空中花粉是植被类型、气候状况与大气环境质量的指示物之一。空中花粉雨的取样实验对化石花粉谱的解释和古植被的恢复具有指导意义,因此空中花粉分析成为第四纪孢粉学和古生态学研究的有效辅助手段。本研究在新疆天山中段北坡进行了短期的空中花粉监测(2001年7月~2006年7月),目的是获得局域性和区域性植被所散发的花粉类型及其数量信息,并探讨花粉数量的植被与气候指示意义、花粉传播以及受环境条件影响的程度,同时分析和检测花粉浓度受气候条件的影响情况。空中花粉的收集使用Cour-type捕捉器,分别在3个不同海拔梯度进行5年连续的空中花粉收集(夏秋季节每周一个样品,冬春季节每两周一个样品)。取样地点分别是新疆天山中段北部的天池气象观测站(43°53'58.38"N, 88°07'15.75"E , 海拔1942.5m)、中科院阜康荒漠生态系统定位站(44°17'27.41"N, 87°55'52.65"E, 海拔477m)和准噶尔盆地古尔班通古特沙漠南缘的北沙窝试验站(44°22'40.74"N, 87°55'9.74"E, 海拔443m),取样点的植被带跨度天山北坡的森林草原到低海拔的典型荒漠。 光学显微镜鉴定的空中花粉主要有43种类型(24科30属),而蕨类、苔藓与真菌孢子数量较少。孢粉类型多样性指数为2.45(Shannon Index)。显然,从样品中所鉴定的空中花粉种类与所观察的植物种类有很大的差距。花粉类型与数量在三个不同海拔梯度的取样点上也有明显差异。天池取样点的花粉类型较丰富,与天山北坡垂直带植被分布以及丰富的植被成分相对应,花粉的优势种类为雪岭云杉(Picea schrenkiana)、蒿属(Artemisia)、藜科(Chenopodiaceae)、柽柳科(Tamaricaceae)和麻黄属(Ephedra),高代表性的外源荒漠花粉类型“削弱了”本地花粉类型的代表性;阜康与北沙窝取样点的花粉类型主要来自荒漠成分的藜科、蒿属、麻黄属、蒺藜科(Zygophyllaceae, 白刺属Nitraria为主)、柽柳科(柽柳属为主)、蓼科(Polygonaceae, 沙拐枣属Calligonum为主)等耐旱植物。花粉数量与植被数量并非简单的线性关系。空中和表土花粉的数量分析表明,藜科、蒿属、麻黄属花粉具有超代表性;雪岭云杉花粉的代表性适中;禾本科(Gramineae)、榆属(Ulmus)、白刺属、柽柳科、沙拐枣属花粉代表性较差。 5年平均花粉浓度大小依次是阜康(81.668 grains/100m3)>天池(51.726 grains/100m3)>北沙窝(45.7685 grains/100m3)。蒿属花粉的峰值出现在秋季,并且出现双峰现象(可能与不同蒿类花期分布不均有关),而其它木本和草本类植物花粉浓度的峰值为单峰,具有明显的季节性,春夏之交开花的桦木属、榆属等类型较为明显。藜科花粉的峰值出现在夏季,麻黄和柽柳的峰值出现在盛夏,桦木属、柳属Salix、雪岭云杉等花粉浓度峰值出现在初夏,与其花期物候相对应。木本植物花粉以雪岭云杉为主,其次为桦木属(Betula)、榆属、柳属、胡颓子科(Elaeagnaceae)花粉,柳属花粉百分比在三个取样点分配比较均匀,桦木属花粉在空间分布上差异明显,与植物分布数量相关。胡颓子科的花粉出现时间短,数量也较少。榆属花粉出现的季节性明显,代表性也较差。因此解释地层中出现类似的低代表性和高代表性的花粉类型时,需要注意其百分比数量的适当校正。木本植物花粉百分比随海拔高度变化成正比关系,对植被的指示性较好。雪岭云杉的年平均花粉浓度在天池取样点明显高于其它两个取样点,5年的平均花粉浓度和百分比与海拔高度呈正比。天池取样点的百分比最高为28.85%,平均为21.15%,而北沙窝与阜康两地5年平均百分比含量分别为0.69%和1.57%,这种时空变化规律与植被数量的关系密切。藜科花粉百分比在阜康和北沙窝取样点占绝对优势,5年的平均花粉浓度与海拔高度呈反比。蒿属花粉年平均浓度与海拔高度关系微弱(R2=0.04,p =0.46),而花粉百分比与海拔高度有显著的线性正比关系(R2=0.72,p<0.0001)。但在低海拔的两个取样点之间差异不明显。 年平均花粉百分比和花粉浓度随海拔梯度而变化,主要花粉类型的变化规律存在差异。暗针叶林雪岭云杉花粉的年平均百分比在低海拔的阜康和北沙窝地区低于3%,在盛花期的6月初,低海拔的取样点也未见超过5%,雪岭云杉花粉的传播在研究区范围内数量分布变化较大,再次表明该类型的花粉在原地沉积效率很高。空中花粉数量能够较好地指示主要的植被带,盛花期内50%以上的雪岭云杉花粉含量则指示了雪岭云杉森林植被带,而高含量的蒿属植物花粉指示了天山山前冲积平原上的蒿类荒漠,相反,高含量的藜科花粉代表了低海拔地区的荒漠。雪岭云杉花粉百分比与浓度均与海拔高度呈显著的线性正比关系,表明雪岭云杉花粉传播的局限性。主要的非木本植物藜科花粉的浓度与海拔高度成反比,蒿属则相反,表明藜科与蒿属花粉在研究区具有很好的植被与气候指示意义。 花粉绝对数量、百分比、浓度具有明显的年际变化,其原因与气候状况的变化有关,年平均花粉浓度、主要类型的花粉浓度与气候参数(降水量、平均温度、最高和最低温度、相对湿度等)的相关分析表明,降水量和湿度与花粉浓度呈负相关程度较高,与其它气候参数则呈显著的正相关。干旱指数、花粉比值、花粉百分比(AP,NAP)与七月份平均气温,年平均降水量的相关系数(Pearson correlation),Ar/Ch与七月平均温度在0.01水平上呈显著的正相关(R2=0.894), 与年平均降水量在0.01水平上呈显著的负相关(R2=0.874)。AP花粉百分比与七月份平均温度呈显著的负相关(R2=0.71, p<0.0001),与年降水量呈正相关(R2=0.43, p=0.01),而NAP花粉正相反。利用短期监测的空中花粉雨数据计算得到的花粉比值以及干旱指数在研究区具有明确的指示意义,尤其是干旱指数、Ar/Ch、Ar/Ep,这些都可为表土花粉、植被与气候关系模型的建立提供重要的参考信息。 气候参数在很大程度上影响大气花粉的浓度,而风速和风向对空中花粉的传播、散布影响较大,花粉的来源与传播受风向和风速的控制。不同取样点的空中花粉数量差异受地形条件影响很明显,这种差异也可利用地形空气动力学特征的差异来解释,天池监测站因“逆坡”(upslope)的气流促进了低海拔的花粉流向高海拔传播。 研究区的空中花粉数量信息是研究表土花粉、植被和气候关系以及第四纪古生态学的重要基础。本研究提高了对研究区空中孢粉的类型及其传播规律的认识和理解,增加了解空中花粉受环境影响的主要因素及其影响程度,对地层孢粉类型鉴定和花粉数据的解释提出指导性建议,对本地区的古植被与古环境研究增加现代孢粉学的参考依据和信息,并可为建立区域性的现代孢粉-植被-气候关系模型提供可靠依据。 同时,本报告也讨论了存在问题和不足,并提出了今后工作的可能完善方案。
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森林作为陆地生态系统的主体,在全球陆地碳循环中起着决定性作用。实测和模型研究均表明北半球的森林是重要的大气CO2汇,在缓解全球碳收支失衡中发挥着关键作用。过去几十年北半球所经历的显著气候变化,已经很大地改变了陆地生态系统的碳平衡状况。随着未来100年气候变化继续增大,对未来气候变化下森林生态系统碳平衡的预测研究就尤为重要。 北京山区森林属于典型的暖温带森林生态系统,前人对本区森林的植被特征、生态系统结构和功能、养分循环以及长期动态变化等都进行了深入的研究。然而长期的人类活动已使本区原生的地带性植被破坏殆尽。因此,对该区域森林生态系统碳平衡的模拟研究可以帮助我们认识其生态系统碳平衡变化特点及未来气候变化对其潜在的影响。 本研究采用LPJ-GUESS植被动态机理模型,利用IPCC于2000年发布的《排放情景特别报告》(SRES)的A2和B2两种情景下不同气候模式对华北地区未来100年温度和降水预测的平均值以及相应大气CO2浓度变化情景进行驱动,模拟北京山区未来100年暖温带森林生态系统的净初级生产力和碳平衡,尽可能真实地反映未来百年的变化趋势。通过比较当前和未来气候情景下北京山区以辽东栎(Quercus liaotungensis)为优势种的落叶阔叶林、以白桦(Betula platyphylla)为主的落叶阔叶林和油松(Pinus tabulaeformis)为主的针阔混交林三种典型暖温带森林生态系统的碳平衡差异,了解未来北京山区这三种暖温带森林生态系统的碳源汇功能,认识气候变化和大气CO2浓度升高对净初级生产力(Net primary productivity, NPP)、净生态系统碳交换(Net ecosystem exchange, NEE)、土壤异养呼吸(Heterotrophic respiration, Rh)和碳储量(Carbon biomass, C biomass)的影响,以及不同生态系统碳平衡对气候变化响应的异质性。 结果表明,未来100年两种气候情景下三种森林生态系统的NPP和Rh均增加,并且A2情景下增加的程度更大。由于三种生态系统树种组成的不同,未来气候情景下各自NPP和Rh增加的比例不同,导致三者NEE的变化也相异:100年后辽东栎林由碳汇转变为弱碳源,白桦林仍保持为碳汇但功能减弱,油松林成为一个更大的碳汇。三种森林生态系统的碳生物量在未来气候情景下均增大,21世纪末与20世纪末相比:辽东栎林在A2情景下碳生物量增加的比例为27.6%,大于B2情景下的19.3%;白桦林和油松林在B2情景下碳生物量增加的比例分别为34.2%和52.2%,大于A2情景下的30.8%和28.4%。各森林类型碳平衡状况不同,原因除气候因素外,主要是由于树种组成的差异所导致。SRES A2和B2两种气候情景相比,相对较低的排放情景(B2)下,生态系统有更高的碳储量。
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能量流动、物质循环和信息传递共同构成了生态系统三大功能。能量是维持生态系统功能与过程的动力,Jordan(1971)认为能量比干物质更能反映出群落对自然资源(特别是太阳能)的利用情况。Long(1934)率先用热值来表示植物所含能量的多少,此后关于植物热值的研究工作逐渐展开。我国的植物热值研究始于20世纪80年代(杨福囤等,1983;祖元刚等,1986),主要集中在海南(林益明等,2000)、福建(林益明等,2004)、广东(任海等,1999)和浙江(郝朝运等,2006)等热带亚热带植物群落以及内蒙古羊草草原(鲍雅静等,2003),但有关黄土高原地区的植被热值研究尚未见报道。子午岭林区是黄土高原主要的天然次生林区,其植被主要是在弃耕地基础上逐渐恢复起来的(邹厚远等,2002)。由于自然因素与人为因素的影响,子午岭的植被在恢复时间上存在较大差异,形成了处于不同演替阶段的各种植被群落类型。本研究选取的6个代表性植被群落分别是沙棘(Hippophae rhamnoides)群落、山杨(Populus davidiana)群落、白桦(Betula platyphylla)群落、油松(Pinus tabulaeform...
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研究人工林凋落叶分解对土壤性质的影响,为防止土壤退化、增加土壤肥力提供理论指导。【方法】采集四川岷江流域上游的4种(连香树(Cercidiphyllum japonicum)、云南松(Pinus yunnanensis)、糙皮桦(Betula1 utilis)和云杉(Picea asperata))林木凋落叶及林地土壤样品,通过对当年凋落叶进行240 d室内分解培养试验,探讨不同凋落叶在分解过程中对土壤性质的影响。【结果】云杉和云南松凋落叶分解使土壤pH值降低,糙皮桦和连香树凋落叶分解使土壤pH值增加;4种凋落叶分解过程中,土壤有机质和全氮含量,土壤微生物量C、N以及4种土壤酶(蔗糖酶、过氧化氢酶、脲酶和蛋白酶)活性均有所增加。【结论】土壤有机质、全N、微生物量、酶活性增加的幅度与凋落叶分解速率及养分释放率有密切关系,凋落叶分解的越快,土壤状况改善的越明显。
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在大气CO2升高和氮(N)沉降增加等全球变化背景下,N元素对生态系统碳(C)、N元素周转过程的影响开始引起越来越多的关注。作为陆地生态系统C库重要组成部分的森林土壤,也逐渐成为研究的重点之一。 本文选择长白山地区典型森林生态系统表层土壤和凋落物,利用人工施N,在实验室控制条件下,模拟N沉降对森林土壤表层C、N元素周转过程影响。旨在从微观上揭示N沉降对凋落物和土壤表层腐殖质分解过程中C、N元素周转过程和土壤C库的影响,主要结论如下: (1) 外源N输入加快了凋落物早期的分解。阔叶树种(岳桦 Betula ermanii、蒙古栎 Quercus mongolica、椴树 Tilia amurensis)的凋落物的分解速度明显快于针叶树种(红松 Pinus koraiensis、鱼鳞云杉 Picea jezoensis)。凋落物的木质素含量是控制其分解速度的主导因子。 (2) N添加对凋落物可溶性有机C(DOC)淋失没有显著影响。DOC淋失主要受凋落物基本性质的控制。阔叶树种的凋落物DOC淋失量明显高于针叶树种。 (3) 不同植被下的土壤性质和C、N周转过程有较大的差异,岳桦林下土壤的微生物生物量和N矿化速率都显著高于暗针叶林,而土壤的C矿化量却低于暗针叶林。岳桦林土壤的DOC和DON淋失量也高于暗针叶林。 (4) N添加显著降低了森林表层土壤的呼吸速率。外加氮对土壤DOC淋失的影响存在一个平衡点,过高的N输入可能加快土壤中DOC的分解速度,降低DOC的淋失量。 研究结果表明,尽管长白山森林生态系统还没有达到“N饱和”,但不断升高的N沉降水平将对长白山典型森林生态系统土壤的C、N元素的周转过程产生较大的影响。但要全面评估N沉降对长白山地区森林土壤C库的影响,还需在野外进行长期定位研究。
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探讨全球气候变化的生物学和生态学效应是当今生态学中的热点,研究大气CO2浓度升高对植物-昆虫相互作用关系的影响具有重要的理论和实践意义。本文使用开顶式气室(Open-top chamber,OTC)在野外条件下研究了CO2浓度升高对三种树木(小青杨、白桦和蒙古栎)叶片化学成分含量的影响,以及树木叶片品质变化对一种广食性森林昆虫(舞毒蛾)幼虫取食、生长发育和取食偏嗜性的影响。得出如下结果:(1)CO2浓度升高对3个受试树种叶片中的营养成分及次生代谢物含量均有显著影响,总体表现为氮含量降低,而碳氮比、非结构性碳水化合物、总酚和缩合丹宁含量增加。叶片中的化学成分含量可随时间发生显著变化,不同树种、甚至同一树种不同冠层高度的叶片对CO2浓度升高的响应强度也是不同的。叶片的干物质含量和比叶重对CO2浓度升高的响应不显著。(2)室内非选择性取食实验、室内选择性取食实验以及上树取食饲养方式下的多龄期取食实验,均发现高浓度CO2处理组内舞毒蛾幼虫的生长发育受到显著抑制。但对四龄舞毒蛾幼虫所进行的短期生物测定并未发现不同CO2浓度处理下幼虫的生长发育速率、对食物的取食率和转化率等昆虫营养指标存在显著差异。(3)叶片品质的降低是导致舞毒蛾幼虫生长发育受抑制的主要原因。但是总体上,CO2浓度升高导致的叶片品质变化并未显著影响幼虫的取食率和取食量。(4)舞毒蛾幼虫对不同叶片种类表现出清晰的取食选择性,这种选择性在其幼龄期就可表现出来。幼虫对小青杨上层叶片有最显著的偏嗜性,对蒙古栎下层叶片有最明显的拒食性。但是CO2浓度升高导致的叶片品质变化对舞毒蛾幼虫的取食选择性和寄主偏嗜行为并未产生显著影响。(5)检测出高浓度CO2处理组内舞毒蛾幼虫虫粪中含有浓度更高的植物次生代谢物质(总酚和缩合单宁),这很可能是昆虫整体生长发育受抑制的重要原因之一。
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近地表面多年冻土对寒区生态系统的植被覆盖、水文条件、土地利用和工程建设具有重要影响,随着气候变化研究的广泛开展,区域冻土环境的变化也成为学者关心的重要议题。中国东北的多年冻土处于欧亚大陆多年冻土带的南缘,多年冻土不如以北地区发育,是十分脆弱的多年冻土。然而,多年冻土在东北寒区生态系统中却起着重要的作用。若东北多年冻土发生退缩,则有可能加速落叶松北移和湿地退缩的过程,也会对C的释放产生重要影响。因而探明现实气候条件下东北区域多年冻土的影响因子和发育状况以及未来气候条件下多年冻土的退缩趋势,将有助于促进东北寒区生态系统的冻土和其它学科研究,同时也可为寒区开发建设提供有意义的参考。 本研究从分析东北多年冻土的主要影响因子——气候、地形和土壤条件等入手,准确地掌握了多年冻土的发育状况,并以此为根据进行了景观尺度上多年冻土分布信息的提取和融深信息的研究。同时,在区域尺度上对多年冻土的现实分布和未来气候条件下多年冻土的可能分布状况进行了探讨。最终得到以下重要结论: (1)冻结数对东北多年冻土分布具有重要的指示作用 冻结数模型具有明确的物理意义,可以指示多年冻土的发生状况。研究中,利用地形、纬度等因子,结合气温和降水数据模拟了现实气候条件下东北地区的冻结数值;并依据冻结数模型的区划标准对东北多年冻土进行分区。结果表明,冻结数在指示多年冻土分布时具有重要作用。 (2)土壤含水量、地形坡度和群落因子对多年冻土具有重要影响 以大兴安岭呼中国家级自然保护区为例,调查了该区多年冻土活动层厚度,并利用多重对比分析和相关分析的统计方法,对多年冻土活动层的影响因子进行了分析。结果表明,多年冻土活动层厚度与多个环境因子之间存在着复杂的关系。其中,土壤表层含水量与活动层厚度具有极显著的负相关关系(P<0.001),其相关系数在0.90以上,说明含水量越高,活动层厚度越浅。地形坡度和活动层厚度的相关性也达到显著水平(相关系数为0.321,P=0.006),表明坡度越陡,活动层厚度越大。几乎每个样带的海拔与活动层厚度都有显著的相关性,但在整体研究区域内海拔与活动层厚度不存在相关性。这说明活动层厚度的变异仅在本研究的样带尺度上具有规律性,而在稍大尺度上这种规律性就消失了。对于不同的群落活动层厚度的多重对比分析表明,群落的差异对活动层厚度也有明显的影响,其中狭叶杜香-泥炭藓群落(Larix gmelini-Ledum palustre var. anqustum-sphagnum magellanicum)更有利于多年冻土的保存。 (3)景观尺度上的多年冻土分布状况 在景观尺度上,以呼中国家级自然保护区为研究区,应用神经网络方法,同时以土地覆盖、等效纬度、坡向和土壤湿度多种影响因子为数据源,对多年冻土分布信息进行提取。结果表明,考虑土地覆盖、等效纬度和土壤湿度的数据源组合可以获得高精度最高的多年冻土分布信息,分类精度可以达到89.0%,多年冻土面积占研究区面积百分比达到46.71%,为780.1 km2。 (4)景观尺度上多年冻土的融深状况 研究考虑了包括植被和等效纬度两个影响活动层厚度的重要因子,并将Stefan公式进行变形,简化为包含热量条件的等效纬度因子和植被条件的C因子的函数关系。最后应用该函数关系模拟了呼中自然保护区活动层厚度空间分布,模拟结果的精度为87.25%。在模拟结果中,面积和所占比例最大的活动层厚度为70-80 cm间的活动层厚度,所占面积达到341.4 km2 ,占整个研究区面积的20.43%。而面积最小的活动层厚度为30-40 cm间的活动层厚度,面积为0.02 km2 。通过群落与活动层厚度的空间分布对比发现,呼中自然保护区占最大比例的活动层(70-80 cm)所对应的植物群落主要为落叶松-丛桦-笃斯-藓类群落(Larix gmelini-Betula ovalifolia-Vaccini uliginosum-moss)。说明呼中自然保护区冻土湿地植被主要以该群落类型为主,演替处于中间阶段。 (5)区域尺度上多年冻土的分布状况 利用证据权重法,以可能影响多年冻土分布的气候、地形和土壤等因子作为数据源,对研究区在现实气候条件下的多年冻土分布进行预测,获得了多年冻土在现实气候条件下的分布概率等信息。结果表明,当分布概率大于0.17时,划分出的多年冻土的精度最高,为78.71 %。此时,多年冻土面积为2.03×104 km2 ,约占研究区总面积的1.76%。 (6)东北多年冻土分布对气候变化的响应 利用空间代时间的方法和Kappa指数,对证据权重法在预测未来气候变化条件下多年冻土分布的准确性进行了验证,结果表明,证据权重法预测气候变化条件下多年冻土的分布状况是可行的。 在CGCM3模拟的三种气候模式下,多年冻土在2050年和2100年都将发生明显的退缩。2050年,SRES A1、SRES A2和SRES B1三种气候情景下多年冻土的面积分别为786.38 km2,705.94 km2和1 028.81 km2。与现实气候下多年冻土的面积2.03×104 km2相比,多年冻土分别退缩了96.13%,96.53%和94.94%。而2100年的模拟结果表明,三种气候情景模式下,多年冻土已经全部退化。 (7)气候变化条件下东北多年冻土的分区变化 研究将冻结数等值图与2000年中国东北冻土分区图进行叠加,计算了不同多年冻土亚区的边界对应的冻结数值,建立了利用冻结数进行中国东北多年冻土分区的标准。根据冻结数指标确定的新的中国东北冻土分区与原中国东北冻土分区进行Kappa指数认证。结果表明,冻结数分区标准更适用于中国东北多年冻土的区划。 利用新的冻结数分区标准对CGCM3模拟的三种气候情景模式下的气候变化数据进行区划表明,三种气候模式下东北多年冻土区在21世纪都会有非常明显的退缩。2050年时冻土区缩减了37.7%-42.6%,2100年时缩减了62.5%-74.0%。同时,研究结果显示,东北多年冻土区域的退缩不仅发生在多年冻土区的南界,同时多年冻土的中心退缩也较为明显,即大片连续多年冻土亚区和大片连续—岛状多年冻土亚区的退缩最为剧烈。2050年时,三种气候情景下,大片连续多年冻土亚区将退缩88.8%以上;2100年时,SRES A2模式下,大片连续多年冻土亚区将完全消失。
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由于森林生态系统的复杂性,过去常用统计回归模型模拟它的各种动态现象。但这样的模型不能揭示森林生态系统的内在规律,故可称之为“黑箱”模型。随着对森林生态系统认识的加深,以及电子计算机技术的广泛应用,用来模仿森林生态系统内在结构与功能规律的各种计算机模型得到了极大发展,从而产生一些不同“灰化”程度的森林动态“仿真”模型。本文建立的阔叶红松林生长与演替计算模型(DOPIDE)就属于这样的“灰箱”模型。KOPIDE(for KOrean PIne'DEciduous mixed forests)是在JABOWA(Botkin, 1972)和FORET(Shugart等,1977)两个模型的基础上建立的,它们都可称为GAP模型。KOPIDE模型共涉及八个树种:红松(Pinus koraiensis)、水曲柳(Fraxinus mandshurica)、紫椴(Tilia amurensis)、春榆(Ulmus japonica)、蒙古柞(Quercus mongolica)、色木(Acer mono)、白桦(Betula platyphylla)、以及山杨(Populus davidiana)。建立该模型的理论基础是森林演替的林窗(CAP)动态理论,它以一年的步长模拟了样地里每株树木的整个生长发育过成(即更新、生长和枯死)。虽然KOPIDE是JABOWA和FORET的直接效仿模型,但它的模拟针对性较强,在模型的结构上有了很大改进。它的运行结果表明,KOPIDE模型在树种生物学特性的描写上、在阔叶红松林动态规律的模拟上、以及在择伐生长的预测上都有较好的可靠性,可用来揭示阔叶红松林长时期的更新、生长和演替规律与特征,也可模拟阔叶红松林的各种经营方式,为决策人提供预测性结果,以供参考。经过KOPIDE模型的反复运行与调试,发现各树种在更新、枯死与林窗大小的关系上很不相同。根据这种现象,可将树种划分为四类:1、在较大的林窗下更新,枯死后不产生大林窗;2、更新不需要林窗,死亡后产生的林窗较大;3、更新需要林窗条件,枯死后不产生林窗;4、不在林窗下更新,死亡后也不产生林窗。尽管这样的建有点绝对化和简单化,但从中可以得出几类树种之间复杂的相互关系网。由此把复杂森林生态系统大范围的动态变化过程视为组成其许多同质或异质小林地单元的动态相嵌。林地单元的同质性,决定了它们动态变化的同步性,从而导致整个林分动态的突性或不稳定性;而林地单元的异质性是决定森林生态系统稳定性的关键性因素。这从另一角度为在东北东部山地和发展阔叶红松林提供了较有说服力的理论基础。
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随着工业化的发展,大气中二氧化碳的浓度(CO2)预测从现在的平均350μmol·mol-1升高到2030年的570μmol·mol-1,其增温作用将持续多个世纪。植被在大气二氧化碳减排以及调控区域水热状况过程中起重要作用,而其机理过程目前十分不清楚。本实验应用自控、封闭、独立生长室,研究了CO2浓度和温度升高对红桦根、茎、叶和枝可溶性蛋白含量和分配的影响,从蛋白水平上来解释川西亚地区的建群种-红桦对CO2升高和温度升高及其交互作用的响应规律,为全球气候变暖川西亚高山的植被保护和恢复提供理论依据。研究结果表明: 1. CO2浓度升高增加了可溶性蛋白的总量,改变了可溶性蛋白分配模式,即,可溶性蛋白分配到根的比例增加,分配到茎、枝、叶的比例减少。可能意味:在CO2浓度升高条件下,红桦根系的生长和营养物质吸收功能将会增强。 2. CO2浓度升高增加了根和茎的清蛋白含量,降低了叶片的清蛋白含量,叶片的球蛋白含量、醇溶蛋白含量和谷蛋白含量均增加。表明CO2浓度升高增加了清蛋白在根中积累,球蛋白、醇溶蛋白和谷蛋白大量在叶片中积累;前人研究所指出的CO2浓度升高使植物叶片可溶性蛋白的含量降低可能仅仅是由于清蛋白含量的降低造成的。 3. 温度升高使红桦幼苗整株所含可溶性蛋白总量增加,但可溶性蛋白总量的分配因红桦幼苗器官的不同而异。温度升高下根、茎、叶和枝的分配量分别占总可溶性蛋白的27.74%、35.57%、23.00%、13.68%,即茎>根>叶>枝。对照的根茎叶枝的分配量分别占总可溶性蛋白的21.01%、41.41%、23.08%、14.50%,即茎>叶>根>枝。表明温度升高使可溶性蛋白分配到根的比例增加,有利于根的可溶性蛋白的积累,增强了根吸收水分和矿质营养的能力,从而有利于根系的生长。 4. 温度升高处理下清蛋白和球蛋白在根中含量升高,在茎、叶和枝中含量下降,但没有达到显著水平;醇溶蛋白在根和叶中含量显著增加;谷蛋白在茎中的含量显著降低。表明温度升高增加清蛋白和球蛋白在红桦幼苗根部的积累,也有利于根和叶醇溶蛋白的积累,但不利于谷蛋白在茎的积累;温度升高条件下叶片可溶性蛋白升高是醇溶蛋白在叶片中积累的结果。 5. CO2浓度和温度同时升高条件下红桦幼苗的可溶性蛋白总量增加很少,只有分配到茎的可溶性蛋白比例增加,并且对可溶性蛋白分配规律没有影响。CO2和温度同时升高下红桦幼苗枝的可溶性蛋白含量的降低是可溶性蛋白总量的降低而不是碳水化合物稀释的结果,并且CO2和温度同时升高对红桦幼苗的生长没有明显的促进作用。 6. CO2和温度同时升高处理对可溶性蛋白含量有显著影响。清蛋白含量在根、茎、叶和枝中均降低,球蛋白含量在根中显著降低,醇溶蛋白含量在根、茎、叶和枝中均降低,谷蛋白含量在根中显著降低。表明CO2浓度和温度同时升高对根的影响显著,即降低了根的可溶性蛋白含量,可能导致根的吸收能力下降。 7. 因此,CO2和温度同时升高对可溶性蛋白影响不能简单地通过CO2和温度单因子影响机理来解释。 It is well known that atmospheric CO2 concentration and temperature are increasing as a consequence of human activities. Atmospheric CO2 concentration are predicted to increase from 350μmol·mol-1 now to 570μmol·mol-1 2030. And temperature will continue to increase for several centuries as a result of CO2 enrichment. Vegetation play a key role in reducing atmospheric CO2 and adapting and controlling warter and energy process in a certain region, while the underlying mechanism are not clear, yet. Betula albo-sinensis, as the dominating tree species of subalpine dark coniferous forest in west Sichuan province, play an important role in determing structure and function of forest ecosystem. In our study, effects of elevated atmospheric CO2 concentration (ambient±350±25μmol·mol-1), increased temperature (ambient±2.0±0.5℃) and their combination on contents and allocation of soluble protein were studied in independent and enclosed-top chamber system under high-frigid conditions. Chambers with ambient CO2 concentration and temperature are taken as control. The results are as the following, 1) Elevated atmospheric CO2 increased the accumulation of total weight of soluble protein in whole plant and changed allocation of soluble protein in red birch by increasing its allocation to roots and reducing its allocation to stem. This caused much more accumulation of soluble protein in roots which might help to prompt growth, development and nutrient absorption ability of roots. 2) Treatment EC increased content of albumin in roots and stems, reduced the content of albumin in leaves, and increased the content of globulin, promalin and glutenin in leaves. That is to say EC increased the accumulation of albumin in roots and accumulation of globulin, promalin and glutenin in leaves. The reduced soluble protein contents in plant leaves by EC, as reported by former researchers, are mainly resulted from the reduced content of albumin in leaves. 3) Elevated temperature increased the total of soluble proteins, but its allocation was dependent on organs. In treatment ET, roots, stems, leaves and branches take 27.74%, 35.57%, 23.00% and 13.68% of total weight of soluble protein. In treatment CK, roots, stems, leaves and branches take 21.01%, 41.41%, 23.08% and 14.50%. Elevated temperature changed allocation of soluble proteins in that it stimulated soluble proteins accumulation in roots and improved the uptake of water in roots. 4) Treatment ET increased the content of albumin and globulin in roots, and reduced the content of albumin and globulin in stems, leaves and branches. The content of promalin in roots and leaves was increased significantly, and the content of glutenin in stems was reduced significant. This suggested that ET stimulated the accumulation of albumin and globulin in roots and accumulation of promalin in leaves and roots; that treatment ET increased content of soluble protein in leaves was mainly resulted from the increased promalin content in leaves. 5) Regarding treatment ETC, the total of weight of soluble proteins increased, but not significantly; but increased in stems. So the combination of elevated atmospheric CO2 and temperature had not changed the allocation of soluble proteins in red birch seedling and reduced soluble proteins in branches were not the result of increased carbohydrate. 6) Treatment ETC reduced the content of albumin and promalin in roots, stems, leaves and branches, reduced the content of globulin and glutenin in roots significantly. That is to say elevated atmospheric CO2 and temperature reduced the content of soluble proteins in roots significantly which might help to prompt growth, development and nutrient absorption ability of roots. 7) The effects of elevated atmospheric CO2 and temperature on soluble protein cannot be simply interpreted through their mechanism that obtained when they were imposed on plant separately.
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在青藏高原东部的亚高山针叶林区,如何尽快恢复这一生态脆弱地区的植被,改变生态环境恶化的趋势,是一个十分重要的课题。光一直被认为是植物种间相互替代,尤其是森林演替过程中植物相互替代或植被恢复中的关键环境要素之一。植物能否适应林冠下或林窗中异质的、或多变的光照条件,对其在林中的生存、分布、更新以及森林动态都是非常重要的。 本文以青藏高原东部亚高山针叶林的主要森林类型——岷江冷杉林群落的几种树苗为研究对象,采用实验生态学、生理及生物化学等方法,通过模拟针叶林不同大小林窗内光照强度的变化,在中国科学院茂县生态站内采用遮荫处理设置6个光照梯度(100、55、40、25、15与7%全光照),来研究具有不同喜光特性的植物对光强的响应与适应机制,其研究结果可为揭示亚高山针叶林的演替规律、以及人工林下幼苗的存活与定居提供科学依据,也能为苗木的生产与管理提供科学指导,尤其是对针阔树种在不同光强下的响应与适应的比较研究,能为如何将阔叶树种整合到人工针叶林中提供新的思路。 光强对植物生长的影响 光强对植物的生长具有重要作用,不同植物在各自适宜的光强梯度下才能生长良好。通过一个野外盆栽实验,来研究不同光强对植物生长的影响(第三章)。主要研究结果如下,低光强下植物株高/茎生物量增加,说明植物会将生物量更多用于高生长,以便有效地拦截光资源;在强光下,植物将生物量更多地向根部分配,使得植物在强光下能够吸收更多的水分,而避免干旱胁迫。 在第一个生长季节,以相对生长速率(RGR)表示,红桦和青榨槭在100%全光照下RGR最大,粗枝云杉在55%最大,岷江冷杉在25-40%下较好;然而,在第二个生长季节,2种阔叶树的相对生长速率(RGR)的适宜光强则变为25-55%,云杉为55-100%,而冷杉为25-100%。可见,从第一年到第二年,2种阔叶树苗更适宜在部分荫蔽的条件下生长;而2种针叶树苗对光的需求则逐渐增加,这可能是增加对根生物量相对投资的结果,因为以这种方式,强光下生长的针叶树幼苗更能保持其内部水分平衡,其生长不会因干旱胁迫而受到严重影响。另外,严重遮荫会引起冷杉幼苗死亡。 植物对光强的生理适应 植物可以通过自身形态和生理特征的调整,来发展不同的光能利用策略从而能够在林中共存。通过一个野外盆栽实验,研究了不同光强下生长的几种树苗的生理特征(第四章)对不同光强的响应与适应。结果显示:强光下,粗枝云杉和红桦的光合能力增加,而岷江冷杉和青榨槭在中度遮荫(25-55%)的条件下光合能力最大。植物叶氮和叶绿素含量增高,而光补偿点和暗呼吸速率降低,这些都是植物对低光环境的适应性反应;而强光下植物叶片和栅栏组织变厚,是对强光的一种保护性反应。 植物对光的可塑性反应 不同植物会表现出对光适应有利的生理和形态可塑性反应。本文对第三章、第四章的实验数据进行可塑性指数分析,来研究植物对光强的表型可塑性反应(第五章)。结果显示,生理特征调整是植物对不同光环境的主要适应途径。红桦和青榨槭的可塑性指数平均值要大于粗枝云杉和岷江冷杉,充分表明这2种阔叶树在生理和形态上较强的可塑性更有利于对光环境的适应,而具有比耐荫树种更强的适应能力。另外,2种针叶树相比,云杉的适应性更强。本研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。 植物的光抑制与防御 当植物叶片吸收了过多光能,会发生光抑制现象。植物对光抑制的敏感性及防御能力对其生长具有重要意义。本文通过两个野外盆栽实验,研究了生长在强光下(第六章)和变化光强下(第八章)植物的光抑制现象及其防御策略。结果表明,在强光下或从遮荫状态转入强光下,植物都会发生光抑制,其对光抑制的敏感性与植物的耐荫性(或喜光)和演替状态有密切联系。长期生长在强光下的植物受到光抑制是可恢复的,而当处于荫蔽环境的植物突然暴露于强光下时,受到的光抑制不能完全恢复,可能是(部分)光合机构受到破坏的缘故。粗枝云杉和青榨槭防御光抑制伤害的能力较强,热耗散是其防御光抑制的主要途径。长期的强光作用能使岷江冷杉和红桦发生严重光抑制,甚至光伤害,而红桦能够通过“凋落老叶,萌发新叶”的途径来适应新的强光环境。 How to restore the vegetation of subalpine coniferous forest in eastern Qinghai-Tibet Plateau, and change the trend of ecological deterioration is a very important issue. Acclimation of tree seedlings to different and varing light environment affects to a great extent the successful regeneration and establishment of subalpine coniferous forests in southwestern China’s montane forest areas, because the ability to respond to such changing resource are commonly assumed to be critical to plant success, and have a growth advantage than others. In this paper, several species seedlings in Abies faxoniana community were chosed to study the response and adaptation to light intensity and the interspecific differences of adaptability in six shaded sheds (100, 55, 40, 25, 15 and 7% of full sunlight) in the Maoxian Ecological Station of Chinese Academy of Sciences. Our results could provide a strong theoretical evidence for understanding the forest succession laws of subalpine coniferous forests, and the survival and settlement of seedlings under plantations, and provide scientific direction for the production and management of seedlings, especially the comparative studies of the acclimation to light between the conifer and broadleaf trees could provide new ideas for how to integrate the broad-leaved trees into the artificial coniferous forest. Growth under different light intensity Light intensity plays an important role on plant growth. One field experiments was conducted to study the growth of tree seedlings of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii under different light intensities. The results showed that plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources, while biomass greater allocation to the roots, could make plants under high light environment absorb more water, and avoid drought stress. During the first growing season, the relative growth rates (RGRs) of Betula albo-sinensis and Acer davidii had the greatest values under the 100% of full light, for 55% of Picea asperata, and for 25-40% of Abies faxoniana. However, in the second growing season the the relative growth rates of the two broad-leaved trees changed and were appropriate for 25-55% of full light, for 55-100% of spruce, and for 25-100% of fir. Thus, from the first year to the second year, two broad-leaved seedlings maybe more suitable to partly shading environment, and two coniferous seedlings would have an increase in light demand, which may be an increased root biomass investment. Because in this way, seedlings grown under high light could better maintain their internal water balance, and thus its growth would not be seriously affected by drought stress. In addition, serious shading would cause fir seedlings to die. Acclimation of physiology to light Plants could coexist in forest ecosystem by forming different strategies of light use. One field experiments was conducted to study the acclimation of tree seedlings to different light intensity of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii. The results showed that the photosynthetic capacity of Picea asperata and Betula albo-sinensis exhibited a general tendency of increase with more light availability; but for Abies faxoniana and Acer davidii seedlings, their highest values of the same parameters were found under intermediate light regime (i.e. 25-55% of PFD relative to full sunlight). Plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources. Leaf nitrogen and chlorophyll content increased, while dark respiration rate and light compensation points decreased, all of which were adaptive response to the low light environment. On the contrary, plants under high light environment had the thicken leaves and palisade tissue, which was a protective response to high light. Phenotypic plasticity to light Phenotypic plasticity can be exhibited in morphological and physiological processes. Physiological characteristical adjustment is the main for plant adaptation to different light environment.The means of plasticity indexes for Betula albo-sinensis and Acer davidii seelings were greater than Picea asperata and Abies faxoniana, amplied that the two broad-leaved trees were much more adaptable to the environment. In addition, spruce had the higher adaptablity than fir. The findings supported the hypothesis that the ecological characteristics of the species determined the biological status and its biological habitat selection. Photoinhibition and photoprotection to light Compared with conifer, broad-leaved trees could better change leaf morphology and adjust biomass allocation to adapt to changing light environment. However, excess light can photoinhibit photosynthesis and may lead to photooxidative destruction of the photosynthetic appatus. Two field experiments were conducted to study the photoinhibition of photosynthesis. The results showed that when plants grown under high light environment or plants transferred from low to high irradiance, the four tree seedlings would undergo a period of photoinhibition. In four species, photoinhibited leaves could recover to initial photosynthetic rates when they were long-term planted under high light environment. However, when plants were suddenly exposed to high irradiance, this photoinhibition could not be reversible, may be the photosynthesis apparatus were (or partly) photooxidatively destructed.
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辐射传输研究是贯穿森林生态系统的纽带,太阳辐射为植物的生长发育提供光合能量、适宜的环境温度以及发育信息。一方面,气候变化使到达地面辐射能的质和量发生变化,影响到植被的生长发育,改变森林的结构,而森林结构的变化又会影响林冠内辐射能的分配和质量,这些变化会进一步影响到林下土壤温度,改变森林根系活性以及土壤营养转化的效率;连锁反应的结果有可能会使森林生态系统的生产力发生变化,改变碳素和氮素源库的调节方向,从而反馈影响地球气候系统。另一方面,人类作为生态系统的成员,必然需要森林生态系统为其提供更多的原材料和更好的生态服务功能,如何实现这些目标,就需要人类适度调整干预方式和频度,达到预期的目的。本文在建立适合于川西亚高山森林的叶面积测量技术、光照辐射模型和土壤温度变化模型的基础上,对川西亚高山地带森林生态系统的辐射传输特征进行了分析,并从森林结构的角度探讨了林分内的辐射分布以及对土壤温度的影响。主要成果如下: 1. 提出了一种照相法测量叶面积的方法。通过对摆放在平面上的叶片照相,利用投影变化,把非正射图像转化为正射图像,然后经过计算机图像处理得到每一片叶片的面积、周长、长度、宽度等信息。这种方法可使用户以任意方向和距离拍摄处于平面上的叶片,能同时处理大量的叶片,适于野外离体或活体叶片测量。叶片面积分辨率可调,分辨率可以与常用的激光叶面积仪相近甚至更高,而且叶片图像可以存档查询。 2. 提出一种模拟林内光照变化的模型。利用林冠半球照片,记录视点以上半球内的林冠构件空间分布,作为林冠子模型;天空辐射子模型采用国际照明委员会(CIE)的标准晴天和阴天以及插值模型。该模型能够模拟林下某一位点处的实时光斑变化。 3. 提出一种土壤温度变化模型。把土壤视为具有容量和阻力性质的结构,利用电阻和电容器件构建土壤能量分布模型。外界太阳辐射能经过植被以及其它一些能量分配器后进入土壤,其中有一部分转化为土壤势能,即土壤温度。土壤温度的变化类似于电池的充放电过程。在已知模型参数的情况下,可以从太阳辐射计算土壤温度的变化。在模型参数未知的情况下,通过输入和输出值推算模型的参数,而模型参数中的时间常数与土壤组成和含水量有关,这样就可以知道土壤水分的变化情况。 4. 从王朗亚高山森林典型样地林分结构的测量获得林地三维结构图、树冠形态、叶面积密度等参数,这些参数输入到Brunner (1998)开发的tRAYci 模型中计算出一段时间内林分任意位置处的光照值。与林下辐射计测量值以及半球照片计算结果的比较,该模型基本上能够满足对林分光环境了解的要求。 5. 从川西亚高山森林生产力的角度,探讨了森林生产力研究的方法以及川西地区的研究历史和成果,发现了其中的一些规律和问题,特别是在叶面积测量上,还没有使用标准的叶面积指数定义。综合来看,川西地区针叶林叶面积指数(单位土地面积上植物冠层总叶面积的一半) 应在4-5 之间。降雨丰富的华西雨屏带是川西地区森林生产力最高的地区,而向西北森林生产力逐渐降低。川西地区云冷杉林森林生产力平均约为600 gDM m-2 a-1,但是根据辐射能计算的潜在生产力则达到1800 gDM m-2 a-1。实际与潜在森林生产力的巨大差异说明其它因子对生产力的影响。 6. 王朗亚高山3 个典型森林林分中,白桦林样地(BF) 林下草本以糙野青茅、牛至、紫菀等喜阳性物种为主,林下透光度较高;冷杉林样地(FF) 林下透光度最低,以喜阴性物种水金凤、蟹甲草、囊瓣芹等为主;而云杉林样地(SF)林分林龄最大,林下透光度介于冷杉林和白桦林之间,草本层仍然以喜阴性物种东方草莓、紫花碎米芥、酢浆草等为主。冷杉林和云杉林的灌木层也很丰富,卫矛属、五加属、茶藨子属、忍冬属植物很丰富,而在白桦林则以栒摘要子属、榛子属、鹅耳枥属等植物为主。藓类植物在云杉林中最丰富,并且形成毯状层,其它两个林分则很稀少。3 个样地林分结构与林下光环境有很强的相关性,从光环境特征可以在一定程度上推测林分的结构。各样地单纯从乔木层材积推算的NPP 排列顺序为BF>FF>SF,与林下辐射透射率和林分年龄的顺序相同,暗示辐射对群落演替过程的驱动作用。 7. 用半球照相法测得BF、FF 和SF 3 个样地的有效叶面积指数以SF 样地最高,BF 最低。如果考虑针叶树叶片在小枝上的丛聚分布,利用北方针叶林的数值进行校正,则SF 样地LAI 显著增加(达到89%),其它样地的LAI 基本不变甚至有所下降。校正后的数值与文献中地面测量的结果较相近,说明在使用半球照相法测量川西亚高山针叶林LAI 时必须加以校正。 8. 在3 个样地中,白桦、岷江冷杉和方枝柏种群为丛聚分布,紫果云杉在FF和SF 样地中基本上为随机分布。3 个物种出现丛聚分布的最短距离约为2m,在最短距离以内则为随机分布。最短距离可能与树冠大小有关,种子传播特征以及对光照的需求状况可能是造成这种分布格局类型的原因。 Radiative transfer plays a key role in forest ecosystems. Solar radiation providesenergy for photosynthesis, appropriate ambient temperature and development informationfor plants. However, quality and quantity of radiation reaching land surface are affected byweather and subsequently influence the growth and development of plants, which in turnchanges the budget of radiation in forest. Soil temperature changes with the variation ofradiation under forest canopy and influences the activity of roots and rate of nutrientturnover. Thus, any changes of radiation will induce chain reactions in the entireecosystem and display in the value of net primary productivity which will possibly shiftthe relationship between carbon source and sink at local or regional scale and feed back tothe global climate system. On the other hand, as a component of ecosystems, humanbeings of course need to demand more materials and better service from ecosystems. Forthese purpose, man must adapt their pattern and frequency of interference to ecosystems.This paper aims to research on the canopy structure, the radiation distribution and theirinfluence on soil temperature from the process of radiative transfer in subalpine forestecosystem of western Sichuan. The main results are: 1 Present a new photogrammetric method for leaf area. The main idea is to convertnon-vertically taken images of planar leaves to orthoimages through projectivetransformation. The resultant images are used to get leaf morphological parametersthrough image processing. This method enables users to take photos at almost anyorientation and distance if only the leaves are placed on same plane, and to processlarge quantity of leaves in a short time, which is suitable for field measurement. Theresolution of leaf area is adjustable to fit for special requirement. 2 A model using hemispherical photos combining with solar tracks and radiation courseis provided to simulate light variation in forest. The hemispherical photos of canopyrecord the real spatial distribution of each element of plants viewed from a point. Skyradiance is simulated with CIE standard clear sky or cloudy sky model. This modelcan be used to simulate real time light variation under canopy. 3 Present a soil temperature model. Soil could be regarded as a body of resistor andcapacitor. Some of the budget of solar radiation in soil body is transformed into soilpotential energy, the soil temperature. Variation of soil temperature is driven by solarradiation, vegetation, soil properties, etc. This model has two parameters, one of whichis time constant and is related to soil water content. The inversed model can be used tosimulate the variation of soil water. 4 By using model tRAYci developed by Brunner (1998), the 3-D distribution of light inthree subalpine forest stands of Wanglang Nature Reserve has been simulated andvalidated with value of radiometers in these stands. This model can basically satisfythe need for understanding light regimes of these stands. 5 Present some principles and questions of NPP (net primary of productivity) researchesin western Sichuan. The standard leaf area index (LAI) defined by Chen and Black(1997) has not been used in this region. Total leaf area and projected leaf area indexare still used in NPP researches which may differ around 1-fold in magnitude. Thestandard LAI which is a half of total leaf area above unit land area should be between4 and 5 for typical subalpine coniferous forest of western Sichuan concluded fromliteratures. The maximum forest NPP occurs in West China rain belt and decreasesnorthwestwards. Average NPP of spruce-fir forest in western Sichuan is about600gDM m-2 a-1, which is below the potential NPP of 1800gDM m-2 a-1 based onmeasured radiation in this region. The significant difference between potential and realNPP suggests that other factors influence the growth of stands. 6 In the three subalpine forest stands of Wanglang Nature Reserve, herbage layer ofAbstractbirch stand (BF) with age of 40 is dominated by heliophytes of Deyeuxia scabrescens,Origanum vulgare, Aster tongoloa etc.. However, both of the other two stands aredominated by shade tolerent species, such as Impatiens noli-tangere, Impatiensdicentra, Cacalia deltophylla and Pternopetalum tanakae etc. in fir stand (FF) withage of 180 and Fragaria orientalis, Cardamine tangutorum and Oxalis corniculata etc.in spruce stand (SF) with age of 330. Shrub species in the latter two stands arerelatively rich, typical dominant genera being Euonymus, Acanthopanax, Ribes andLonicera. Birch stand has relatively sparse shrubs dominated by genera of Cotoneaster,Corylus and Carpinus. Mosses are significant only in spruce stand. The canopystructure controls the light regime of stand, which influence the composition of herblayers beneath the canopy. This light regime-community structure relationship can beused to infer the herb community from canopy structure. The NPP derived from timbervolume of arbor layer of the three stands decreases from BF to SF, which is in thesame order of transmitted total radiation under canopy and age of these stands,suggesting the driving effect of radiation in the succession of community. 7 The highest effective LAI of the three stands obtained by hemispherical photos is inplot SF and lowest in plot BF. After rectification of the clumping effect of leaves onshoot, the real LAI in plot SF increases significantly (89%) and approximate to theaverage LAI of coniferous forest in western Sichuan. Therefore, the LAI obtainedfrom hemispherical photos needs rectification for clumping effect. 8 Spatial distribution pattern for Betula platyphylla, Abies faxoniana and Sabinasaltuaria is clumpy, but Picea purpurea almost random in plot FF and SF. The shortestdistance for clumpy distribution for Betula platyphylla and Sabina saltuaria is 1.5m,and 2m for Abies faxoniana. And random pattern for these trees is exhibited within thisrange which almost coincides with the diameter of crown. Seed dispersalcharacteristics and light requirement may be the reason for different spatial pattern.
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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 ¡æ,并预测在本世纪将上升1.4-5.8 ¡æ。气候变暖对陆地植物和生态系统产生深远影响,并已成为全球变化研究的重要议题。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。森林采伐迹地和人工云杉林下作为目前该区人工造林和森林更新的两种重要生境,二者截然不同的光环境对亚高山针叶林不同物种更新及森林动态有非常重要的影响。 本文以青藏高原东部亚高山针叶林几种主要森林树种为研究对象,采用开顶式增温法(OTCs)模拟气候变暖来研究增温对生长在两种不同光环境下(全光条件和林下低光环境)的几种幼苗早期生长和生理的影响,旨在从更新角度探讨亚高山针叶林生态系统不同树种对气候变暖在形态或生理上的响应差异,其研究结果可在一定程度上为预测气候变暖对亚高山针叶林物种组成和演替动态提供科学依据,同时也可为未来林业生产管理者提供科学指导。 1、与框外对照相比,OTCs 框内微环境发生了一些变化。OTCs 框内与框外对照气温年平均值分别为5.72 ¡æ和5.21 ¡æ,而地表温度年平均值分别为5.34 ¡æ和5.04 ¡æ,OTCs 使气温和地表年平均温度分别提高了0.51 ¡æ和0.34 ¡æ;OTCs框内空气湿度年平均值约高于框外对照,二者分别为90.4 %和85.3 %。 2、增温促进了三种幼苗生长和生物量的积累,但增温效果与幼苗种类及所处的光环境有关。无论在全光或林下低光条件下,增温条件下云杉幼苗株高、地径、分支数、总生物量及组分生物量(根、茎、叶重)都显著地增加;增温仅在全光条件下使红桦幼苗株高、地径、总生物量及组分生物量(根、茎、叶重)等参数显著地增加,而在林下低光条件下增温对幼苗生长和生物量积累的影响效果不明显;冷杉幼苗生长对增温的响应则与红桦幼苗相反,增温仅在林下低光条件下对冷杉幼苗生长和形态的影响才有明显的促进作用。 增温对三种幼苗的生物量分配模式产生了不同的影响,并且这种影响也与幼苗所处的光环境有关。无论在全光或林下低光环境下,增温都促使云杉幼苗将更多的生物量分配到植物地下部分,从而导致幼苗在增温条件下有更高的R/S 比;增温仅在林下低光条件下促使冷杉幼苗将更多的生物量投入到植物叶部,从而使幼苗R/S 比显著地降低;增温在全光条件下对红桦幼苗生物量分配的影响趋势与冷杉幼苗在低光条件下相似,即增温在全光条件下促使红桦幼苗分配更多的生物量到植物同化部分—叶部。 3、增温对亚高山针叶林生态系统中三种幼苗气体交换和生理表现的影响总体表现为正效应(Positive),即增温促进了几种幼苗的生理活动及其表现:(i)无论在全光或林下低光环境下,增温使三种幼苗的光合色素含量都有所增加;(ii)增温在一定程度上提高了三种使幼苗的PSII 光系统效率(Fv/Fm),从而使幼苗具有更强的光合电子传递活性;增温在一定程度使三种幼苗潜在的热耗散能力(NPQ)都有所增强,从而提高幼苗防御光氧化的能力;(iii)从研究结果来看,增温通过增加光合色素含量和表观量子效率等参数而促进幼苗的光合作用过程。总体来说增温对幼苗生理过程的影响效果与幼苗种类及所处的光环境有关,增温仅在全光条件下对红桦幼苗光合过程的影响才有明显的效果,而冷杉幼苗则相反,增温仅在低光条件下才对幼苗的生理过程有显著的影响。 4、增温对三种幼苗的抗氧化酶系统产生了一定的影响。从总体来说,增温使几种幼苗活性氧含量及膜脂过氧化作用降低,从而在一定程度上减轻了该区低温对植物生长的消极影响;增温倾向表明使三种幼苗体内抗氧化酶活性和非酶促作用有所提高,从而有利于维持活性氧代谢平衡。但增温影响效果与幼苗种类所处的光环境及抗氧化酶种类有关,增温对冷杉幼苗抗氧化酶活性的影响仅在林下低光环境下效果明显,而对红桦幼苗抗氧化酶活性的影响仅在全光条件下才有明显的效果。 总之,增温促进了亚高山针叶林生态系统中三种幼苗的生长和生理表现,但幼苗生长和生理对增温的响应随植物种类及所处的光环境不同而变化,这种响应差可能异赋予了不同植物种类在未来气候变暖背景下面对不同环境条件时具有不同的适应力和竞争优势,从而对亚高山针叶林生态系统物种组成和森林动态产生潜在的影响。 Enrichment of atmospheric greenhouse gases resulted from human activities suchas fossil fuel burning and deforestation has increased global mean temperature by 0.6¡æ in the 20th century and is predicted to increase it by 1.4-5.8 ¡æ. The globalwarming will have profound, long-term impacts on terrestrial plants and ecosystems.The ecoologcial consequences arising from global warming have also become thevery important issuses of global change research. The subalpine coniferous forests inthe eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying theeffects of climate warming on terrestrial ecosystems. The light environment differssignificantly between clear-outs and spruce plantations, which is particularlyimportant for plant regeneration and forest dynamics in the subalpine coniferous forests. In this paper, the short-term effects of two levels of air temperature (ambient andwarmed) and light (full light and ca. 10% of full light regimes) on the early growthand physiology of Picea asperata, Abies faxoniana and Betula albo-sinensis seedlingswas determined using open-top chambers (OTCs). The aim of the present study wasto understand the differences between tree species in their responses to experimentalwarming from the perspective of regeneration. Our results could provide insights intothe effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientificdirection for the production and management under future climate change. 1. The OTCs manipulation slightly altered thermal conditions during the growingseason compared with the outside chambers. The annual mean air temperature andsoil surface temperature was 5.72 and 5.34 ¡æ (within the chambers), and 5.21 and5.04 ¡æ (outside the chambers), respectively. The OTCs manipulation increased airtemperature and soil surface temperature by 0.51 and 0.34 ¡æ on average, respectively.Air relative humidity was slightly higher inside the OTCs compared with the controlplots, with 90.4 and 85.3 %, respectively. 2. Warming generally stimulated the growth and biomass accumulation of thethree tree species, but the effects of warming on growth and development variedbetween light conditions and species. Irrespective of light regimes, warmingsignificantly increased plant height, root collar diameter, total biomass, componentbiomass (stem, foliar and root biomass) and the number of branches in P. asperataseedlings; For A. faxoniana seedlings, significant effects of warming on all the tested parameters (plant height, root collar diameter, total biomass, and component biomass) were found only under low light conditions; In contrast, the growth responses of B.albo-sinensis seedlings to warming were found only under full light conditions. Warming had pronounced effects on the pattern of carbon allocation. Irrespectiveof light regimes, the P. asperata seedlings allocated relatively more biomass to rootsin responses to warming, which led to a higher R/S. Significant effects of warming onbiomass allocation were only found for the A. faxoniana seedlings grown under lowlight conditions, with significantly increased in leaf mass ratio (LMR) and decreasedin R/S in responses to warming manipulation. The carbon allocation responses of B.albo-sinensis seedling to warming under full light conditions were similar with theresponse of A. faxoniana seedlings grown under low light conditions. Warmingsignificantly decreased root mass ratio (RMR), and increased leaf mass ratio (LMR)and shoot/root biomass ratio (S/R) for the B. albo-sinensis seedlings grown under full light conditions. 3. Warming generally had a beneficial effect on physiological processes of dominant tree species in subalpine coniferous forest ecosystems: (i) Warming markedincreased the concentrations of photosynthetic pigments in both tree species, but theeffects of warming on photosynthetic pigments were greater under low lightconditions than under full light conditions for the two conifers; (ii) Warming tended toenhance the efficiency of PSII in terms of increase in Fv/Fm, which was related tohigher chloroplast electron transport activity; and enhance non-radiative energydissipation in terms of in increase in NPQ, which may reflect an increased capacity inpreventing photooxidation; (iii) Warming may enhance photosynthesis and advancephysiological activity in plants by increasing photosynthetic pigment concentration,the efficiency of PSII and apparent quantum yield (Φ) etc. From the results, theeffects of warming on seedlings’ physiological performance varied between lightenvironment and species. The effects of warming on photosynthesis performance of B.albo-sinesis seedlings were pronounced only under full light conditions, while thephysiological responses of A. faxoniana seedlings to warming were found only underthe 60-year plantation. These results provided further support for the observationsabove on growth responses of seedlings to warming. 4. Warming had marked effects on antioxidative systems of the three seedlings.Warming generally decreased H2O2 accumulation and the rate of O2- production, andalleviated degree of lipid peroxidation in terms of decreased MDA content, whichalleviated to some extent the negative effects of low temperature on the plant growthand development in this region; Warming tended to increase the activities ofantioxidative enzymes and stimulate the role of non-enzymatic AOS scavenging,which helped to create an balance in maintaining AOS metabolites for the threeseedlings. Nevertheless, the effects of warming on antioxidative defense systems werepronounced only under the 60-year plantation for the A. faxoniana seedlings. Incontrast, the marked effects of warming on antioxidative defense systems for the B.albo-sinesis seedlings were found only under the full light conditions. In sum, warming is considered to be generally positive in terms of growth andphysiological process. However, the responses of growth and physiology performanceto warming manipulation varied between species and light regimes. Competitive and adaptive relationships between tree species may be altered as a result of responsedifferences to warming manipulation, which is one mechanism by which globalwarming will alter species composition and forest dynamics of subalpine coniferousforest ecosystems under future climate change.
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黄龙世界自然遗产地岷江冷杉林(Abies faxoniana)生境类型多样,群落结构复杂,群落植物种类组成多样性丰富。揭示不同生境的生物多样性及其差异是认识生物多样性格局、形成及维持机制的前提和进行多样性保育的基础。本文采用样方法对黄龙钙化滩生境、阴坡非钙化生境及半阳坡非钙化生境的岷江冷杉原始林植物群落结构及植物多样性进行了研究。结果表明: 黄龙岷江冷杉林具有明显的复层异龄结构,垂直结构明显,乔木、灌木、草本、苔藓层次分明。共发现高等植物386 种,其中维管植物46 科103 属163 种,苔藓植38 科83 属物223 种。各层片结构及物种组成如下: (1)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境分别发现乔木18 种、13种、8 种。乔木层均可分为两个亚层,第一亚层优势种均为岷江冷杉,第二亚层主要为岷江冷杉异龄树或其它大高位芽物种。钙化滩生境第一亚层除优势种岷江冷杉外混生有巴山冷杉(Abies fargesii)、粗枝云杉(Picea asperata)以及阔叶树种白桦(Betula platyphylla)等,第二亚层主要为岷江冷杉异龄树;阴坡非钙化生境第一亚层除优势种岷江冷杉外间有巴山冷杉和白桦,第二亚层物种主要为川滇长尾槭(Acer caudatum var. prattii);半阳坡非钙化生境第一亚层除优势种岷江冷杉外混生有巴山冷杉,第二亚层主要为岷江冷杉异龄树。依乔木层优势种的差异,钙化滩生境及半阳坡非钙化生境为岷江冷杉纯林,阴坡非钙化生境为岷江冷杉-川滇长尾槭混交林。不同生境乔木层郁闭度、乔木密度、树高结构、直径结构均存在差异。 (2)钙化滩生境发现灌木41 种,平均盖度为18.49±1.72(%),平均高度为52.12±4.45(cm),优势种为直穗小檗(Berberis dasystachya);阴坡非钙化生境发现灌木30 种,平均盖度为29.33±2.56 (%),平均高度为119.55±8.01 (cm),优势种为箭竹 (Fargesia spathacea) 、唐古特忍冬(Lonicera tangutica) 和袋花忍冬(Lonicera saccata);半阳坡非钙化生境发现灌木29 种,平均盖度为31.35±1.93 (%),平均高度为107.55±4.24 (cm),优势种为箭竹(Fargesia spathacea)。不同生境灌木层结构和物种组成多样性差异显著,钙化滩生境的灌木盖度、高度总体上较非钙化的坡地生境低, 钙化滩生境灌木以小型叶的落叶灌木为主,沟两侧非钙化的坡地生境上则发育了丰富箭竹。 (3)钙化滩生境发现草本46 种,平均盖度为7.18±0.79 (%),平均高度为5.04±0.26(cm),以山酢浆草(Oxalis griffithii)为优势种;阴坡非钙化生境发现草本物种71 种,平均盖度达29.04±2.31(%),平均高度为9.08±0.52(cm),以钝叶楼梯草(Elatostema obtusum)、山酢浆草为优势种;半阳坡非钙化生境草本物种50 种,平均盖度为以8.79±0.82(%),平均高度为7.67±0.43 (cm),以扇叶铁线蕨(Adiantum flabellulatum)、双花堇菜(Viola biflora)、华中蛾眉蕨(Lunathyrium shennongense)、山酢浆草为优势种。阴坡非钙化生境草本层片发育良好,多样性最为丰富,盖度和物种丰富度均显著高于钙化滩生境和半阳坡非钙化生境。 (4)钙化滩生境发现苔藓物种140 种,平均盖度达84.25±1.30 (%),以仰叶星塔藓(Hylocomiastrum umbratum) 等大型藓类为优势种;阴坡非钙化生境发现苔藓物种115 种,平均盖度为79.29±1.64 (%),以刺叶提灯藓(Mnium spinosum)、大羽藓(Thuidium cymbifolium)、毛尖燕尾藓(Bryhnia trichomitra)等个体较小的物种为优势种;半阳坡非钙化生境发现苔藓物种91 种,平均盖度为60.64±1.93 (%),也以刺叶提灯藓为优势种。 (5)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境的物种数分别为234 种、221 种、175 种。乔木层的Shannon-Wiener 指数分别为0.75 ±0.12、1.87±0.12、1.78±0.07(灌木层,0.44±0.08、1.71± 0.15、2.49±0.06;草本层,0.33±0.13、1.31±0.15 、2.15±0.08; 苔藓层1.30±0.11、2.08±0.04、1.73±0.11,);Pielou 均匀度指数分别为0.45±0.05、0.29±0.06、0.28±0.08(灌木层,0.75±0.03、0.68±0.05、0.52±0.06;草本层,0.68±0.02、0.77±0.02、0.74±0.02;苔藓层,0.40±0.03、0.63±0.02、0.52±0.03);Simpson's 优势度指数分别为0.63±0.06、0.78±0.04、0.83±0.07(灌木层,0.21±0.03、0.28±0.05、0.45±0.06;草本层,0.25±0.02、0.12±0.01、0.17±0.01;苔藓层,0.45±0.04、0.18±0.01、0.31±0.04)。三种生境间乔木层、草本层的Sorenson 群落相似性系数较低, 灌木层、苔藓层的的Sorenson 群落相似性系数较高。 综上所述,黄龙岷江冷杉林的群落结构、植物多样性在三种生境间存在差异性,这将意味着我们在进行黄龙世界自然遗产地的森林经营管理时要较多地关注岷江冷山林群落在不同生境中的差异性。 There were multiplex habitat types, complicated community structure and abundant species composition in the Huanglong World Natural Heritage Site. Uncovering the differences of biodiversity among different habitats was a precondition to understand the distribution, formation and sustaining mechanism of the biodiversity, and the foundation of biodiversity conservation. In the present study, using plenty of quadrants, we investigated the community structure and the biodiversity of the primitive Abies faxoniana forest in different habitats (travertine bottomland, semi-sunny-slope non-calcified habitat and shady-slope non-calcified habitat) in the Huanglong World Natural Heritage Site. The main results are as follows: All the primitive Abies faxoniana forests in the three habitats were uneven-aged with obvious vertical structure including tree layer, shrub layer, herb layer and bryophyte layer. A total of 386 higher plants including 163 vascular plant species (103 generic, 46 families) and 223 bryophyte species (83 generic, 38 families) were investigated. The structure and species composition of each layer are as follows: (1) There were 18, 13 and 8 tree species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. The tree layers in all habitats can be divided into two clear sub-layers. The upper tree layers were dominated by Abies faxoniana, and the lower tree layers were dominated by uneven-aged Abies faxoniana or other phanerophytes species. There were Abies fargesii , Picea asperata and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in travertine bottomland, and the lower tree layers were dominated by uneven-aged Abies faxoniana; There were Abies fargesii and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in shady-slope non-calcified habitat, and the lower tree layers were dominated by Acer caudatum var. prattii; There was Abies fargesii besides the dominated species (Abies faxoniana) in the upper tree layer semi-sunny-slope non-calcified habitat, and the lower tree layers were dominated by uneven-aged Abies faxoniana. According to composition percentage of dominate species in tree layer, both the forest in travertine bottomland and in semi-sunny-slope non-calcified habitat could be ranked as pure forest, and the forest in shady-slope non-calcified habitat could be ranked as mingled forest. There were significant differences in crown density, plant density, height structure and diameter structure among the three habitats. (2) A total of 41 shrub species (average coverage 18.49±1.72%; average height 52.12±4.45 ㎝)were found in travertine bottomland, and the dominate species was Berberis dasystachya; A total of 30 shrub species (average coverage 29.33±2.56 %;average height 119.55±8.01 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Fargesia spathacea, Lonicera tangutica and Lonicera saccata. A total of 29 shrub species (average coverage 31.35±1.93%; average height 107.55±4.24 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Fargesia spathacea. There were significant differences in structure and species diversity of the shrub layers among the three habitats. The coverage and height of shrub had lower value in travertine bottomland than in two non-calcified habitats. Moreover, travertine bottomland was dominated by deciduous shrub species with microphyll and non-calcified habitats developed abundant Fargesia spathacea species. (3) A total of 46 herb species (average coverage 7.18±0.79%;average height 5.04±0.26 ㎝)were found in travertine bottomland, and the dominate species was Oxalis griffithii; A total of 71 herb species (average coverage 29.04±2.31%;average height 9.08±0.52 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Elatostema obtusum and Oxalis griffithii. A total of 50 herb species (average coverage 8.79±0.82%;average height 7.67±0.43 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Adiantum flabellulatum, Viola biflora, Lunathyrium shennongense and Oxalis griffithii. Herb layers developed well in shady-slope non-calcified habitat and had the higher species richness and coverage than travertine bottomland and semi-sunny-slope non-calcified habitat. (4) A total of 140 bryophyte species (average coverage 84.25±1.30%)were found in travertine bottomland, and the dominate species was big bryophyte species such as Hylocomiastrum umbratum and so on; A total of 115 bryophyte species (average coverage 79.29±1.64%)were found in shady-slope non-calcified habitat, and the dominate species was small bryophyte species such as Mnium spinosum, Thuidium cymbifolium, Bryhnia trichomitra and so on. A total of 91 bryophyte species (average coverage 60.64±1.93%) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Mnium spinosum. (5) There were 234, 221 and 175 plant species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. Shannon-Wiener index of the tree layer was 0.75 ±0.12, 1.87±0.12 and 1.78±0.07 (the shrub layer, 0.44±0.08, 1.71± 0.15 and 2.49±0.06; the herb layer, 0.33±0.13, 1.31±0.15 and 2.15±0.08; the bryophyte layer, 1.30±0.11, 2.08±0.04 and 1.73±0.11.) for the three habitats, respectively; Pielou index of the tree layer was 0.45±0.05, 0.29±0.06 and 0.28±0.08 (the shrub layer, 0.75±0.03, 0.68±0.05 and 0.52±0.06; the herb layer, 0.68±0.02, 0.77±0.02 and 0.74±0.02; the bryophyte layer, 0.40±0.03, 0.63±0.02 and 0.52±0.03.) for the three habitats, respectively. Simpson's index of the tree layer was 0.63±0.06, 0.78±0.04 and 0.83±0.07 (the shrub layer, 0.21±0.03、0.28±0.05、0.45±0.06; the herb layer, 0.25±0.02, 0.12±0.01 and 0.17±0.01; the bryophyte layer, 0.45±0.04, 0.18±0.01 and 0.31±0.04.) for the three habitats, respectively. There were low Sorenson index both in the tree layer and in the herb layer among the three habitats, whereas, high Sorenson index occurred both in the shrub layer and in the bryophyte layer. To sum up, there were differences both in community structure and plant diversity among the three different habitats, which means that we should pay more attention to habitats heterogeneities of the primitive Abies faxoniana forest when we take action to manage the forest in the Huanglong World Natural Heritage Site.
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Knowledge about cumulative effects of forest management alternatives on forest landscape is required to make forest management decision. In this paper,a spatially explicit landscape model,LANDIS,was applied to simulate forest landscape changes in 200 years under four management alternatives (no cutting,clearcutting,selective cutting I and II) in Youhao Forestry Bureau located in Small Khingan Mountains. APACK was used to calculate distribution area of the representative species and species age cohort for six species. The results showed:1) timber harvest decreased area percentage of representative conifer species,Pinus koraiensis,Picea koraiensis and Picea jezoensis,Larix gmelinii to some extent compared to no cutting. The most influencing cutting mode for the area percentage of Tilia amurensis and Quercus mongolica was selective cutting II,followed by selective cutting I and clearcutting. To the contrast,the change of area percentage of Betula phatyphylla was contrary to the management alternatives; 2) As to species age cohort composition,timber harvest significantly changed age structure,that is,it decreased over-matured age cohort of representative species,and increased seedling and middle-age cohort (B. phatyphylla was not included,because its area percentage of over-mature age cohort was the highest under clearcutting than other three scenarios).
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通常认为,随着林木不断接近其海拔分布极限,光合作用产量不断下降,导致碳水化合物供应不足(碳供应限制),或者低温限制了碳投资(生长限制)。植物组织内非结构性碳水化合物(Nonstructural carbohydrates,NSC)的含量反映了植物碳供应与碳吸收的平衡。为了检验"碳供应限制"和"生长抑制"假说,我们对长白山海拔1700~2050m的自然生境下生长的岳桦(Betula ermanii)的叶片和枝条组织的NSC含量进行了比较。结果表明:岳桦叶片的NSC含量随海拔升高变化不显著,枝条的NSC含量随海拔升高显著增加;叶片和枝条中淀粉含量与可溶性总糖含量的比值均随海拔的升高而减小;林线附近的岳桦林木不存在碳水化合物供应不足的问题,这在一定程度上表明生长限制导致长白山岳桦林线的形成。