增强UV-B辐射对暖温带落叶阔叶林下土庄绣线菊水分生理和氮素利用及形态特性的影响

在中国北方大部分地区，水分始终是影响植物生长和分布的最主要限制性因子之一，植物在其生长期经常遭受水分胁迫。不仅如此，随着大气同温层中臭氧浓度的减少，过量的有害紫外辐射（主要是UV-B，280nm-320nm）将穿透大气层达到地球表面。随着全球变化的加剧，这些地区的植物将不可避免地受UV-B和水分胁迫的共同作用。 本实验是在北京东灵山暖温带森林生态系统中，选择常见灌丛土庄绣线菊(Spiraea pubescens)，建立UV-B控制实验。连续三个生长季每天增补9.4kJ•m-2的辐射剂量，模拟臭氧衰减17%时近地表面UV-B辐射的增强。本实验的目的是观测在野外环境下，长时间人工增强UV-B辐射对土庄绣线菊水分生理、氮素利用以及形态特征的影响。具体对以下指标进行测定：叶片的气孔导度、碳同位素比率（δ13 C）、叶含水量、叶面积、水分利用效率（WUE）、叶全氮含量、叶氮素再吸收率。 实验结果表明，增强UV-B辐射显著减少了土庄绣线菊的叶面积（50.1%），提高了叶片全氮含量(102%)，处理植株的氮素再吸收率比对照植株高出50.9%。同时，UV-B辐射还在一定程度上（尽管统计显示不显著）降低了气孔导度(16.1%)、胞间CO2浓度与大气CO2浓度之比(Ci/Ca) (4.0%)、提高了碳同位素比率（δ13 C）(20.5‰)、叶含水量(3.1%)及比叶重（SLW）(5.2%)，从而导致水分利用效率（WUE）的增加(4.1%)，植物的抗旱能力增强。值得注意的是，深层土壤（30-40cm）含水量变化会影响气孔导度、δ13 C、WUE对紫外辐射的响应程度：在土壤干旱的季节（6月和9月），气孔导度、δ13 C、WUE这些指标处理和对照的差异很小，但是当土壤水分充足时（7月和8月），处理和对照的差异就较为显著。另外，随着实验处理时间的延长UV-B的效应变得不显著。相关分析表明，UV-B辐射降低了土壤含水量（30-40cm）与土庄绣线菊叶含水量、δ13 C、Ci/Ca、气孔导度的相关系数，增强了WUE与土壤含水量的相关性，这也许是由于UV-B辐射增强了WUE对土壤水分变化的敏感性。本研究的结果表明UV-B辐射对土庄绣线菊的形态和生长有显著的影响，但对主要水分生理指标影响不显著。

拉萨河流域亚高山落叶阔叶灌丛的植被生态学及其合理利用与保护的初步研究

拉萨河流域的亚高山落叶阔叶灌丛，以前尚无专门报导。本文对该类灌丛下的土壤、植物区系成份、生活型作了分析、统计，并应用TWINSPAN和DCA方法把野外调查的样地划分为七个群落类型，分别描述它们的群落特征：在将亚高山落叶阔叶灌丛与同其在垂直带中上下相接的其它灌丛分别对比后，对其植被地理学问题作了探讨，并进一步确定了这类灌丛的性质。研究表明．亚高山落叶阔叶灌丛因处于向森林区的过渡地带上，而与之具有一定的联系;在植被分布上表现为拉萨地区的相对偏湿气候与垂直带的产物，因而不能认为是本地区的“高原地带性”植被。关于该类灌丛的来源方面，初步认为包含有原生以及次生类型，还有待于进一步研究。最后，探讨了该类灌丛及群落中经济植物的利用与保护问题：其中着重于充分利用当地的自然条件，增加植被覆盖，进行植被改造。

暖温带落叶阔叶混交林能量生态学研究

本文是首篇研究中国暖温带落叶阔叶混交林能量生态学的论文。在文中，笔者以详实的第一手资料从能量环境、能量流动、能量组合以及能量平衡几个方面，全面、系统地阐述、分析了辽东栎林——这一暖温带落叶阔叶混交林典型自然群落代表的能量生态学特征。 在能量环境一章中，笔者从能量流动，能量平衡的角度出发重点研究了辽东栎群落的辐射能量环境特征。笔者以1991-1993年的观测资料为基础，从乔木、灌木和草本三个层次分析了生长季总辐射、散射辐射、直射辐射、反射辐射、净辐射、先合有效辐射、透射辐射、吸收辐射以及乔木层和灌木层反射率的季节动态和日进程特征，并从天文因子、气象因子和群落自身发育特征几方面解释分析了辐射能量环境的这种时空动态特征，同时，分析了这种变化特征对群落能量流动、分配和平衡过程可能产生的影响。 另外，笔者也对群落湿度和风速环境的时空动态特征进行了分析。 在能量流动一章，笔者以1992-1993年的野外实验资料为依据，沿季节动态、月际变化和日进程的时间轴，从群落、乔木层、灌木层、草本层以及各乔、灌木种群的空间尺度详细分析、阐述了太阳辐射能在森林群落内的流动和转化特征，并从能量环境和群落发育的角度解释分析了能量在群落内的这种时空分布和转化特征。所讨论的能流对象包括群落、乔、灌、草各层及各乔、灌木种群的总能流固定量、叶片呼吸耗能量、剩余能流固定量以及沿枯枝落叶流出的能流量。 与分析能流过程同步，笔者从上述的时、空尺度分别以生长季内太阳总辐射和光合有效辐射为基础计算、分析了森林群落的光能转化率特征。 在这一章的最后，笔者概述性地介绍了辽东栎群落的能量平衡特征 在第四章，笔者从能值的角度出发，以能量密度为标准讨论了能量沿群落各层及各乔、灌木种群的积累、分配和组合特征，并讨论了能量流动和光能转化率与热值和能量密度的关系。 辽东栎群落能量生态学的研究不但为了解暖温带落叶阔叶林生态系统的结构和功能，为恢复和重建退化的森林生态系统提供了丰富详实的理论信息，而且，也为山区人工林优化模式的组建提供了理论依据和实践指南。

森林生态系统的养分循环-一个温带落叶阔叶林生态系统的研究

l、有机质和全N在草甸土壤的不同层次之间呈现明显的差异，表现力随着土壤深度的增加，含量逐渐降低。无机N包括NH4+-N和No3--N在土层之闻的差异并没有象全N和有机质那么显著。NH4+-在表面30Cm的土壤中明显高于NO3--N的。全P在各层土壤中的含量差异并不显著，但有效p在三层土壤中表现出明显的差异，表现为随土壤深度的增加而降低。 K、Na、Ca、Mg、Fe和AI等6种金属元素的可交换成分在革甸土壤中的含量除K表现出明显的土层间的差异外，在表层30cm的土壤中并没有明显的随土壤深度变化的趋势。 有机质和全N在东灵山地区主要森林土壤中的含量以0~lOcm土层中的显著高于10~20cm的土层中的。20cm的土层中无机N中也以NH4+-为主，这与草甸土壤中的一致。全p和有效P在两个土层之间的差异并不十分显著，而以0^lOcm土层略高于10～20Cm土层的。 K、Na、Ca、Mg、Fe和AI等6种金属元素的可交换成分在不同土层间的差异以K和Ca较为明显，K和Ca在O—1Ocm的土层中要高于lO-20cm的。 森林土壤中的有机质、全N（包括NH4+-N）、全p（包括有效P)都表现为低于亚高山草甸土壤中的，主要原因是亚高山革甸分布于海拔1800cm至2300m，而主要森林类型则基本分布在海拔120m至1500m，亚高山草甸较为恶劣的气候条件明显不利于有机质的矿化，因而分解速率较低，积聚了较多的有机质．而森林土壤中较低的NH4+-N和有效P则可能是由于森林中旺盛的生命活动，尤其是微生物的活动，而大量被植物和微生物固持，因而，即使在森林土壤中有较高的矿化速率，但仍具有较低的NH4+-N和有效p的含量． 不同类型的森林土壤之间各化学成分并没有发现有明显的变化趋势，即使森林类型差异较大的落叶阔叶林和针叶林之间也并没有发现十分显著的差异，这说明各森林类型土壤之间养分特征的一致性。两种针叶林均为人工林，林龄不超过30，而在这之前推测也是以落叶阔叶林为主，森林土壤在这30年间并没有发生十分明显的变化。 在l100m至I500m之间的各种森林类型的土壤也没有表现出十分明显的与海拔高度的相关性。同一森林土壤在不同时间的采样之间，各化学成分也没有表现出明显的差异。 2、降水引起的树干茎流水分中全N的含量在降水季节的初期的含量略高于大量降水期间的。而在降水季节后期，随着每次降雨量的减少，降水间隔时间的增加，树干茎流中N的含量明显地有很大幅度的升高。NO3-N在树干茎流中的含量也同样受到了降水量的影响，表现在后期也发生显著的升高，但升高的幅度没有全N的大。受到降水量影响的元素还有S。 全p在树干茎流水分中含量随降水时间的变化，在各个树种之间有很大的差异。 其它几种元素的可交换态在茎流水分中的含量以K的含量为最高;其次是S、ca、Mg，AI在所有分析的元素中含量最低。不过，AI在降水初期(6月22日)的取样中，含量在几个树种的茎流水分样品中普遍较高，Ca和Mg也被发现有类似的情形。 穿透雨中全N的含量在降水初期略高于降水中期的，而至降水末期时又略有升高，这种格局与树干茎流水分中的具有一致性。全P、K、Ca、Mg、S．ＡＩ等包括全N及其无视成分与树干茎流中的含量的差异并不显著． 地表径流中的全N含量在去除枯枝落叶层的径流场的变化格局与树干茎流和穿透雨中的较为一致．表明这三者之间存在一定的相关性．但与未去除枯枝落叶层的径流场的径流水分中全N和NO３－Ｎ的含量并不十分一致，全N和NO３－N的含量有时较高． 取自马牙石沟流水堰口的集水区水样中全N和NO３-N的含量均变化在l～3ｍｇ／L之间;而取自南沟流水堰口的集水区水样中的全N和N03-N的含量均在l～2mg／L之间，但在这两者之间并没有十分显著的差异，说明取自两个集水区溪流水分中的全N和N03-N的含量基本一致。 3、通过对三个森林立地乔木层生物量的两次调查（相隔6年），发现尽管以辽东栎为主要优势树种的暖温带落叶阔叶混交林（91- 03）第一次调查的生物量低于辽东栋林（91 - 02）和桦木林（91 - 04)的，但在生长6年后，该立地的乔木层生物量已明显超过桦木林的，其增长速率在三个样地中为最高。 根据两次调查的结果，三个样地中出现的11种乔木树种胸径的增加可以分成两组，第一组包括北京花楸、糠椴、白桦和辽东栎，另一组则由山杨、五角枫、棘皮桦、大叶白蜡、蒙椴、黄花柳和沙涞组成。6年中，第一组树种的胸径增加量在1.O～l.4em之间，而第二组的增量在0.2~O.9cm之间，两者之间具有显著的差异。 常用的生物量预测模型包括两个因子，即胸径(D)和树高(H)。D通常可以实测得到，但H大部分只能估计得到。本研究通过比较6年前后两次树高的实际估测结果，证明用D和H共同建立的模型预测结果可能会产生较大的人为误差，建议改用D的单因子预测模型。 无论是实测结果还是树木年轮分析结果，都表明处于目前径级水平的暖温带地区分布的这几个树种还处在不断生长的过程中，随着年龄的增加，表现为胸径还有很大程度的增加，因为这些树种目前的大径级个体的年平均增加速率大于小径级的． 树木年龄与胸径的相互关系的分析结果表明，树木年龄和胸径可以很好地用饱和模型Y=a-be－ｃｘ来拟合． 4、通过近5年对辽东栎枝条( D<0.5cm)和叶片凋落物分解的观察和分析，发现辽东栎枝条的总失重率达到了43%，分解速率常数k为2.713x10－3／周，而叶的总失重率则可以达到70%，分解速率常数k为6.234xl0-3/周. 枝条分解过程中的有机质含量变化可以分为三种类型，一种是单调上升型的，如蛋白质，其含量从3.5%增加到约6%;-种是单调下降型的，如半纤维素，约从15%降低至7%;另一种则是相对变化不大，如粗纤维（包括纤维素和木质素），粗纤维的含量变化在58％±3%，木质素变化在56%±2.5%，纤维素则变化在1%～3%之间。 用Olson指数方程拟合，结果表明粗纤维、木质素和半纤维素三者的分解以半纤维素为最快，分解速率常数为5.693xl0-3／周，其次是木质素和粗纤维，分别为2.461x10-3/周和2.352xl0-3／周．而蛋白质和纤维素的拟合结果较差或根本无法拟合． 用Olsoｎ指数方程拟合的结果表明，有些元素在凋落物分解过程中可以很好地用该方程拟合，如K，Na、Mn和C，p和N，Mg，Zn，而一些元素如Ca和Cu没有取得很好的拟合效果。 几种元素在叶片凋落物分解过程过程的丢失速率也以K为最快，其它元素的顺序为，e，N，p，Na，Mn，Mg和Ca（Ca的拟合效果较差），而Cu、Zn也不能很好地用Olson指数方程拟合。 5、根据对东灵山地区6年(199l~1997)的气象观测结果，可得到该地区的年平均降水量和总降水量，用1994年分析的降水中养分含量的数据可以近似计算得东灵山地区每年通过降水进入该暖温带落叶阔叶林生态系统的几种养分输入量。同理可以计算降水输入养分在树干茎流和穿透雨中的分配，还可以计算养分从地表径流中的输出。通过对这几个水分循环的主要环节中养分输入或输出量的计算，发现每年通过降水而进入系统的养分，大部分将积累在系统中，而输出份额只占很小的一部分。 K和Mg两种养分元素在降水通过林冠和树干表面时，将产生大量的淋溶，而Ca的淋溶较小．N、P和AI则产生少量的吸附或吸收。对于S来说，淋溶和吸收或吸附作用相当，或两者都很小。 通过生物量的年平均增量和生物量中的养分含量可以计算得到每年积累在生物量中的养分量即存留量，同理可以计算年凋落物中的养分量、归还量及生物系统从土壤中吸收的养分量，每年养分的存留率．通过以上计算，发现该落叶阔叶林对养分的存留率较低，这可能是由于该落叶阔叶林森林生态系统每年具有较高的养分归还量，对于一些元素来说，还具有较高的淋溶量，尤其是K和Mg．

北京山区落叶阔叶林优势种水分生理生态研究

本文研究了北京山区落叶阀叶林优势种一辽东栎、大叶白蜡、北京丁香、核桃楸、山杏和荆条等乔灌木的稳定碳同位素比率，长期水分利用效率、植物蒸腾特性和土壤植物体的水分运动特点，并从植物解剖学的角度研究了这些植物叶片特点和其水分生理生态特性的关系。 对北京山区落叶阔叶林生态系统这几种乔、灌木植物叶片中的碳稳定性同位素比率(δ¹³C值)测定结果显示，这些植物叶片的δ¹³C值受多种因子的影响，具有较大的种间差异及时空异质性。主要表现在不同植物种叶片δ¹³C值不同，其排列顺序为山杏(- 24.75±0.85%。>大叶白蜡(- 25,94±1.52%。)=荆条(- 26.01±1.63%。)=辽东栎(一26.07±1.17%。)=北京丁香(-26.46±0.80c70。)>核桃楸(-28.11±1.52%。);生长初期叶片δ¹³C值较生长末期高，尤以核桃楸和辽东栎表现明显其生长初期和末期的叶片δ¹³C值皆相差达3‰;生境条件，特别是土壤水分含量和土层厚度，对植物叶片的813C值的高低有较大的影响，生长在于旱生境中的植物具有较高的δ¹³C值。另外，即使是同一株植物，叶片δ¹³C值也因其在冠层中所处的位置不同而异，冠层项部叶片的δ¹³C值高于林冠内部的叶片。北京山区落叶阔叶林优势种的长期水分利用效率与种的特点有关，山杏最高(4.950±0．l71mmolC0_2•mol-1H_2O)，核桃楸最低(3.760±0.203mmolC0_2•mol-1H_2O)，大叶白蜡、荆条、辽东栎和北京丁香居中(4.346- 4.530 mmolCO_2•mol-1H_2O)，大部分植物长期水分利用效率在春季（5月）较高，秋季较低，荆条由于物候期的特殊性在其生长季初期较低，而后逐渐增高。核桃楸在不存在水分亏缺情况下，树干液流速率受微气候因子的影响，液流速率的最大值达1600g•hour 左右。树干液流速率的日进程和大气相对湿度、温度的日进程具有相当好的生态学同步性。通过对核桃楸夜晚树干液流的分析可以得出其有根压存在的结论。 植物叶片和枝条中自由水和束缚水含量主要决定于植物种的特性，枝条的年龄、生境特点，特别是土壤水分特点。在落叶阔叶林I(样地2)中植物叶片自由水含量的排列顺序是：北京丁香>核桃楸>大叶白蜡>辽东栎;而杂灌丛（样地1）中植物叶片自由水含量的排列顺序是：核桃楸>大叶白蜡>山杏>荆条>北京丁香>辽东栎，可见群落类型对植物自由水含量影响是很大的，植物束缚水含量与其自由水含量的格局完全相反，荆条、山杏等植物含量高，核桃揪含量低。枝条水分含量有与叶片水分含量相类似的特点。 北京山区落叶阔叶林优势种的水分生理生态学特性和其叶片的特点有很大的关系，首先是植物叶片的特点总是和其种的特性相联系，主要表现在叶的类型、叶片上毛、气孔密度、着生方式等，如荆条叶片上下表面都密被披针形毛，气孔小，核桃楸气孔较大且凸出，大叶白蜡叶片上的气孔凹陷，辽东栎的气孔呈椭圆状，保卫细胞上有许多白色蜡质结晶。有一些种有环状的气孔外缘。生境的变化对叶片的形态特征有影响，在全光照条件下叶片小而犀，而在庇荫条件下叶片大而薄，在扫描电镜下可见全光照条件下北京丁香叶片基本无毛，庇荫条件下则有短微毛，全光条件下荆条叶片上毛有小乳头状凸起，庇荫条件下没有。本文所研究的植物种气孔都着生在叶片的下表面，气孔密度的大小排列顺序是：辽东栎>山杏>北京丁香>核桃楸>大叶白蜡。经方差分析显示种闷气孔密度存在极显著性差异。对所研究植物的气孔导度和环境因子、叶片解剖特点进行线性回归分析，得到了总体的和各个种的回归模型，结果表明光照强度、气孔密度等对气孔导度影响显著，但因种的不同相互之间存在差别。 植物的蒸腾速率受多因子的影响，主要有种的特点、微气候因子（光照强度、大气相对湿度、叶面温度、叶室温度等）和土壤水分特点。植物的蒸腾速率日进程和微气候因子日进程有相当好的生态学同步性。对辽东栎的蒸腾速率和光照强度的研究发现二者有很好的线性关系。这些植物蒸腾速率都表现出一定规律的日进程和季节进程，大多数植物蒸腾速率在一天中有数次波动，最高峰一般出现在中午12时之前;在整个生长季中，6月底至8月初的蒸腾速率高于其他月份。

落叶栎树在中国的地理替代分布及其气候制约

栎属（Quercus L.）按落叶习性可自然分为落叶类型和常绿类型，我国落叶栎类共有20个种和9个变种。落叶栎是栎属中较为进化的一个类群，源于横断山区和云贵高原;除新疆外，全国各省都有落叶栎的天然分布，一些亲缘关系密切的树种之间呈现出较为明显的地理替代分布格局。本研究的目的在于：（1）应用BIOCLIM模型模拟预测落叶栎类植物的潜在分布区，分析其目前的分布格局以及下一步的发展趋势;（2）分析造成落叶栎树地理替代分布格局的主导气候因子，探讨气候因子对不同落叶栎树种地理分布格局的制约作用。 本文以16个在中国具有成片天然分布区的落叶栎树种（包括变种）为研究对象，利用已核对的标本数据以及13个栅格化环境变量图层（分辨率为1km×1km），按照分类（全国广布型、南方广布型、南方狭域型和北方狭域型）和不分类（全部16种）两种处理方式，通过BIOCLIM模型模拟得出了它们的潜在核心分布区和潜在边缘分布区。在运行模型之前，除必选的海拔高程图层外，采用了主成分分析（PCA）的方法从30个候选的气候变量图层中筛选出对相应落叶栎树种的地理分布格局有较大影响的12个图层作为输入图层。然后，本文通过比较两种处理所得模拟结果的ROC（Receiver Operator Characteristic）曲线下方面积AUC（Area Under the Curve），同时结合文献分析来推测不同落叶栎树种地理分布格局的稳定性及发展趋势。结果表明，在无人类活动干扰且种源传播不受阻碍的情况下，全国广布型和南方广布型落叶栎目前的分布格局在维持稳定的基础上有向周边地区扩展的趋势;南方狭域型和北方狭域型落叶栎的分布格局则基本保持稳定，短期内发生扩散的可能很小。 论文中计算了每个落叶栎树种所在分布范围的气候指标（共11个），以便进行下一步的研究。以蒙古栎（Q. mongolica）、辽东栎（Q. wutaishanica）与槲栎（Q. aliena）、锐齿槲栎（Q. aliena var. acuteserrata）、北京槲栎（Q. aliena var. pekingensis）这两组地理替代系列为研究对象，分别采用独立样本t检验和单因素方差分析的方法，分析了气候因子对其地理替代分布格局的主导作用。结果表明，冬季的低温、较高的气温年较差和大陆度是蒙古栎向东北替代辽东栎的主要原因;槲栎向北被北京槲栎和锐齿槲栎替代的主要原因是生长季高温和冬季高温对其分布的双重制约;除最暖月（7月）最高温外，北京槲栎的各项水热指标与另两种槲栎均存在极显著差异，对冬季低温和较大的年较差的适应可能是限制其向南分布的主要原因。 本研究最后部分的内容是对不同类型落叶栎分布区的气候参数进行的主成分分析。结果表明，生长季温度是制约落叶栎分布的最主要的气候因子;寒冷程度和冬季的低温则对其在大尺度范围的扩散有较大影响;另外，降水、年较差与大陆度对落叶栎的向北分布也起着重要的作用。

SEXUAL DIMORPHISM IN THE SNUB-NOSED LANGURS (COLOBINAE, RHINOPITHECUS)

Sexual dimorphism in the dentition and skeleton of the four extant species of snub-nosed langurs, Rhinopithecus (R.) bieti, R. (R.) brelichi, R. (R.) roxellana and R. (Presbytiscus) avunculus, was studied. The species shared a similar general pattern of sexual dimorphism, but were found to differ in respects that appear to reflect the influence of disparate socioecological and environmental factors. All the species showed marked canine dimorphism, but the very high degree of canine dimorphism in R. bieti appeared to be due to the intensity of intermale competition for mates during a temporally restricted breeding season, and possibly also to the intensity of competition between males for other resources during other times of the year. Sexual dimorphism in the postcranial skeleton of Rhinopithecus species was also most pronounced in R. bieti and may be related to the relatively higher frequency of terrestrial locomotion in males of the species. (C) 1995 Wiley-Liss, Inc.

Sleeping sites of Rhinopithecus bieti at Mt. Fuhe, Yunnan

Data on sleeping site selection were collected for a group of black-and-white snub-nosed monkeys (Rhinopithecus bieti; around 80) at Mt. Fuhe, Yunnan, China (99degrees20'E, 26degrees25'N, about 3,000 m asl) from November 2000 to January 2002. At the site mainly three vegetation types were present in an elevation-ascending order: deciduous broad leaf forest, mixed coniferous and broad leaf forest, and dark coniferous forest. In addition, bamboo forest presented in areas burned in 1958. Sleeping sites (n = 10) were located in the coniferous forest, where trees were the tallest, bottommost branches were the highest, the diameter of crowns was the second largest, and the gradient of the ground was the steepest. Monkeys usually kept quiet during entering and staying at a sleeping site. The site choice and the quietness may be tactics to avoid potential predators. In the coniferous forest, however, monkeys did not sleep in the valley bottom where trees were the largest, but frequently slept in the middle of the slope towards the east/southeast, in the shadow of ridges in three other directions, to avoid strong wind and to access sunshine; in winter-spring, they ranged in a more southern and lower area than in summer-autumn. These may be behavioral strategies to minimize energy stress in the cold habitat. Monkeys often slept in the same sleeping site on consecutive nights, which reflected a reduced pressure of predation probably due to either the effectiveness of anti-predation through sleeping site selection, or the population decline of predators with increasing human activities in the habitat. The group's behavioral responses to interactive and sometimes conflicting traits of the habitat are site-specific and conform to expectations for a temperate zone primate.

子午岭60年辽东栎林种子质量与森林更新

辽东栎是我国暖温带落叶阔叶林地带的优势乔木树种之一。通过26年的定位监测,对子午岭辽东栎林种子质量消减、种子萌发与环境条件、实生苗时空分布动态以及对其森林更新的影响等方面进行了初步探讨。结果表明:辽东栎林在子午岭半阳坡、半阴坡和阴坡3种类型中,平均完好种子占种子总数的26.65%;霉变种子占18.72%;动物取食虫蛀种子在阴坡远高于半阴坡和半阳坡,占到种子总数的26.32%;已发芽的种子占其总数的28.31%,且半阴坡>半阳坡>阴坡。每年均有大量种子生产,但在生境与动物的共同作用下,种子数量和质量受到很大影响,多达73.35%的种子失去生命力,直接影响实生苗的形成;地表覆盖物虽能促进种子的快速发芽,但对成苗却是一个物理障碍,影响是负作用的,主要影响因子是地表覆盖物的厚度和含水量;在辽东栎林下虽有一定的实生幼苗分布,但数量极少,平均密度仅为140~120株/hm2,且不同立地条件差异显著,严重影响森林的天然更新。

KOPIDE：长白山北坡阔叶红松林生长与演替的计算机模型

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。

次生植被土壤碳、氮养分季节动态及凋落物输入与温度的影响

土壤微生物量、可溶性有机碳与氮虽然只占土壤有机碳、氮总量的较小部分，但可以在土壤全碳、氮变化之前反映土壤微小的变化，又直接参与土壤生物化学转化过程，因而在植被恢复过程中，较其它土壤理化性质等能够更好地指示土壤恢复情况。在青藏高原东缘存在大面积的次生人工林替代灌丛或采伐迹地，而关于这些人工林替代后的生态效果和生态过程的评估却十分缺乏，本研究通过评估岷江上游植被恢复重建过程中典型人工替代次生植被凋落物层与土壤碳、氮等养分大小，动态监测土壤微生物生物量、水溶性碳、氮等指标，结合温度与凋落物输入等影响土壤活性有机碳、氮因子的控制试验，系统分析不同人工替代次生植被土壤碳、氮等养分的差异原因，试图寻找低效人工林优化调控与持续管理技术，为区域生态公益林持续管理提供理论和技术依据。主要结论如下： 1. 通过对不同人工替代次生植被凋落物层和土壤碳、氮分析发现，油松和华山松人工林替代次生灌丛后土壤碳、氮含量较灌丛和阔叶人工林低，主要原因可能为凋落物质量(C/N)较差，而引起碳、氮等元素难以归还土壤。进而通过对不同人工替代次生植被凋落物层和土壤微生物生物量、水溶性有机碳、氮等指标的季节性动态模式的分析，发现各次生植被土壤微生物生物量C、N，P以及土壤水溶性碳、氮含量均呈明显季节性动态，呈现秋季明显大于其它季节，冬季最低，在表层土壤最为明显。 2. 油松、华山松人工林凋落物层和土壤水溶性有机碳(WDOC)、土壤水溶性有机氮(WDON)明显低于灌丛和连香树，土壤微生物生物量C、N也以油松和华山松人工林最低，而落叶类植被，如灌丛、连香树和落叶松之间没有明显差异，说明可利用底物的数量和质量差异是影响各次生植被凋落物分解和土壤微生物活性的主要原因。MBC/OC和MBN/ON能较好地指示土壤微生物活性的变化，MBC/OC凋落层总体以灌丛和连香树人工林最高，油松和华山松人工林最低；而土壤中MBC/OC连香树人工最高，华山松人工林最低。说明以油松和华山松为主的人工造林替代乡土阔叶灌丛造成土壤C、N等养分严重匮乏，微生物活性低下是影响其养分周转的主要原因。 3. 从各次生植被凋落物产生看，凋落物年归还量最大的为华山松人工林(5.1×103 kg ha-1)，其次为落叶松人工林(4.8×103 kg ha-1)，阔叶灌丛林地凋落物产生总量(4.4×103 kg ha-1)略大于油松人工林(4.2×103 kg ha-1)，最小的为连香树人工林(3.6×103 kg ha-1)；叶是凋落物的主体，落叶类树种月动态表现为单峰型，高峰主要在10-11月，如落叶松、连香树和灌丛林；常绿的松类月动态不明显，各月基本相同，最为明显地为油松林，华山松人工林略有二个小峰，分别出现在11月和5月。落叶阔叶灌丛的凋落物分解速率大于常绿针叶林，如油松和华山松。结合凋落物的产生量和分解速率，不同树种人工林替代次生阔叶灌丛后，人工油松和华山松林枯落物总贮量和厚度明显大于落叶松人工林、灌丛林和连香树人工林，说明以油松和华山松为主的人工造林替代乡土阔叶灌丛延缓了有机物向土壤的顺利归还，不利于土壤C、N等养分循环。 4. 通过控制地面凋落物和地下根系输入有机物对土壤碳、氮的影响研究发现，(1) 单独去除根系以及根系与地面凋落物同时去除处理1年后对表层(0-10cm)土壤WDOC均没有显著影响，而土壤WDON显著增加，油松人工林土壤微生物生物量C、N显著降低，人工落叶松林没有显著差异，说明油松人工林土壤微生物活性对地下碳输入的依赖大于其它次生植被，而落叶松土壤微生物活性对地下碳输入依赖性较小；去除地面凋落物，明显降低了落叶松人工林土壤WDOC，华山松和连香树土壤WDON均较对照显著减少，油松人工林土壤微生物量C较对照显著减少；双倍增加地面凋落物处理对土壤微生物生物量、WDOC和WDON没有明显地增加，相反，连香树、华山松和油松人工林土壤WDON较对照减少。说明油松人工林微生物活性不仅依赖于地下碳输入，而且对地上有机物输入的依赖性也较大；连香树、落叶松和华山松人工林土壤微生物生物量并没有因地面凋落物的去除减少可能与土壤总有机碳含量及活性均较高有关，而双倍增加地面凋落物反而降低了土壤微生物生物量，说明凋落物覆盖后改变了土壤微气候。 5. 碳矿化累积量与有机碳含量和活性有机碳含量之间存在显著地正相关关系。凋落物碳累积矿化量、矿化速率以连香树最高，油松和华山松人工林次之，落叶阔叶灌丛低于常绿针叶纯林，导致其差异的主要原因可能为凋落物产生的时间动态模式不一样，致使凋落物起始分解时间不一致。而土壤层有机碳矿化速率和矿化量以阔叶落叶灌丛和连香树最高，油松和华山松人工土壤最低，再次证实利用针叶纯林恢复植被阻碍了有机质周转与循环。 6. 凋落物累积矿化量与C/N值呈显著地相关关系，并随着温度的升高而明显增加，而土壤累积矿化量与C/N值没有显著相关关系，说明土壤有机碳质量(C/N)对温度的响应不十分明显。通过双指数模型对不同温度下碳矿化过程进行模拟和计算出活性有机碳与惰性有机碳比例，发现温度升高促进了惰性有机碳向活性有机碳的转化，增加了活性有机碳含量，说明温度升高可促进次生植被凋落物与土壤有机质的分解，进而可影响到林地碳源/汇关系的变化。 综上，通过对不同人工替代次生植被凋落物与土壤C、N大小、以及土壤微生物生物量、水溶性C、N等指标动态变化模式研究，结合温度与凋落物数量输入等影响土壤活性C、N因子的综合分析，以油松和华山松人工纯林对山地植被恢复，延缓或阻碍了有机质周转与循环，造成了土壤肥力退化。对现有低效人工纯林改造，应为地面大量有机物分解创造条件。 Although soil microbial biomass, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) are a small part of total soil organic carbon and nitrogen, they can directly participate in the process of soil biochemical translation and indicate the fine changes before changes of soil total organic carbon and nitrogen occur. So, they are good indexes to indicate soil restoration condition during the process of vegetation rehabilitation. There are large areas of secondary vegetations which substitute for indigenous shrubs in the eastern fringe of Qinghai-Tibet Plateau. However, it is not well known that the ecological effect and process after substitution by different secondary plantations. Based on comparison of soil organic and nitrogen contents in litter layer and soil under different secondary vegetations in upper reaches of Minjiang River, soil microbial biomass, DOC and DON in litter layer and soil were investigated in order to analyze the seasonal dynamic. Combining the effects of temperature, litter input and root exclusion on soil microbial biomass, DOC and DON, we also aim to understand the reason and mechanism of difference in soil carbon and nitrogen contents among different secondary vegetations. The study would contribute to comprehensively understanding C and N cycling processes and provide optimal control and sustainable technology of low-effect plantations in these regions. The results are as follows: (1) Organic carbon and nitrogen in litter layers and soil under different substitution plantations were investigated. The results showed that contents of soil organic carbon and nitrogen were lower in P. tabulaeformis (PT) and P. armandi Franch(PA) than those in native broad-leaf shrub and broad-leaf plantation. The low quality (C/N) of litter in PT and PA plantations caused carbon and nitrogen returning to soil difficultly. Seasonal dynamic of soil microbial carbon (MBC),-nitrogen (MBN),-phosphor (MBP), and WDOC and WDON showed similar pattern, which had the highest values in autumn and the lowest values in winter. (2) WDOC and WDON in litter layers and soil under PT and PA plantations were significantly lower than those in native broad-leaf shrub and Cercidiphyllum japonicum Sieb. et Zucc.(CJ). Soil MBC and MBN were also the lowest, while there were no significant differences among deciduous vegetations, i.e. native broad-leaf shrub, CJ and Larix kaempferi Lamb.(LK) plantation. The results suggested that difference in quantity and quality of available substance was main reason that affected the activity of microbe in soil and litter layer. MBC/OC and MBN/ON were good indexes to indicate the change of soil microbial activity. MBC/OC of litter had the highest value under native broad-leaf shrub and CJ plantation, and had the lowest value in PT and PA plantations, while MBC/OC of soil was the highest under CJ plantation, and was the lowest in PT and PA plantations. These results indicated that PT and PA plantations substituting for native broad-leaf shrub caused deficit of carbon and nitrogen in soil, low microbial activity was a main reason influencing the cycling and turnover of carbon and nitrogen in soil. (3) The annual litter fall production, composition, seasonal dynamic and decomposition of five typical secondary stands in upper reaches of Minjiang River were studied in this paper. The annual litter productions were: PA (5.1×103 kg ha-1), LK(4.8×103 kg ha-1), native broad-leaf shrub (4.4×103 kg ha-1), PT(4.2×103 kg ha-1)，CJ(3.6×103 kg ha-1). The litter production of leaves in five secondary vegetations occupied a higher percentage in the annual total litter production than those of other components. The litterfall was mostly producted in the cool and dry period (October-November) for deciduous vegetations and relatively equably producted in every season for evergreen coniferous vegetations. The decomposition rate of leaf litter in the broad-leaf stand was higher than those in evergreen coniferous stand. Combined with annual litter fall production and decomposition rate of leaf litter, we found that stock and depth of litter layer were significantly larger in PT and PA plantations than those in native broad-leaf shrub, LK and CJ plantations. The results confirmed that PT and PA plantations substituting for native broad-leaf shrub delayed organic matter returning to soil and hindered cycling of carbon and nitrogen again. (4) We explored plant litter removal, double litter addition, root trenching, and combining root trenching and litter removal treatments to examine the effects of above- and belowground carbon inputs on soil microbial biomass, WDOC and WDON in four secondary plantations. During the experimental period from June 2007 to July 2008, 1 year after initiation of the treatments, WDOC in soil did not vary in root trenching, and combining root trenching and litter removal treatments, while WDON in soil significantly increased compared with CK treatment. Root trenching reduced soil MBC and MBN in PT plantation, while MBC and MBN in soil did not vary in LK plantation. The rasults implied that soil microbial activity was more dependent on belowground carbon input in PT plantation than those in other secondary plantations, on the contrary, soil microbial activity in LK plantation was not dependent on belowground carbon input. Plant litter removal significantly decreased soil WDOC in LK plantation, decreased WDON in PA and CJ plantations, and also significantly reduced soil MBC in PT plantation. However, double litter addition did not increase soil microbial biomass, WDOC and WDON, on the contrary, soil WDON in CJ, PA and PT plantations were decreased. These suggested that soil microbial activity was not only dependent on belowground carbon input, but also on aboveground organic material input. Double litter addition could change the microclimate and result in the decrease of soil microbial activity in CJ, PA and PT plantations. (5) We measured carbon mineralization in a 107 days incubation experiment in 5℃,15℃ and 25℃. Carbon cumulative mineralization was positively correlated with organic matter and labile organic carbon in litter layer and soil. Cumulative carbon mineralization and mineralization rate of litter layers in PT and PA plantations were higher than that in native broad-leaf shrub. This difference between native broad-leaf shrub and coniferous plantations in cumulative carbon mineralization and mineralization rate of litter layers could be attributed to the initiating time of decomposition due to the difference in seasonal dynamic of litter fall production between two types of secondary plantations. However, cumulative carbon mineralization and mineralization rate in soil were the highest in native broad-leaf shrub and CJ plantation, and were the lowest in PT and PA plantations. This also confirmed that PT and PA plantations substituting for native broad-leaf shrub hindered the cycling and turnover of organic matter again. (6) Carbon cumulative mineralization was positively correlated with C/N in litter layer and increased with temperature increasing, while carbon cumulative mineralization was not correlated with C/N in soil. This indicated that soil organic matter quality (C/N) was insensitive to temperature. Applying bi-exponential model, we computed the percent of labile and stable carbon in different temperature incubation and found that temperature increasing would accelerate the transform from stable carbon to labile carbon and increase the percentage of labile organic carbon. This illuminated that temperature incraesing could facilitate the decomposition of litter and soil organic matter in secondary vegetations and hence affect the relationship between carbon source and sink.

川西亚高山针叶林主要针叶树种年轮生态学研究

米亚罗地区是四川西部较为典型的亚高山针叶林区域之一。为建立该地区主要针叶树种岷江冷杉、云杉、紫果云杉和红杉的年轮宽度年表资料，了解不同海拔高度岷江冷杉原始林和不同恢复过程的人工针叶林及次生混交林树木径向生长规律，结合样地调查，用生长锥钻取了树木芯样做年轮生态学分析。芯样经过标准化程序固定和打磨抛光后，用WinDENDRO图像分析系统测量年轮宽度序列，用COFECHA程序交叉定年和控制测量数据质量，用ARSTAN程序建立了4个主要针叶树种的地区年表和不同海拔高度岷江冷杉林及人工针叶林和次生混交林针叶树的样地年表。 4个主要针叶树种年轮宽度年表的平均敏感度低于0.2，而其晚材宽度年表都具相对较高的平均敏感度。早材宽度与年轮总宽度标准化年表间的相关系数均在0.9以上；晚材宽度与年轮总宽度标准化年表间的相关系数则种间差异较大，红杉的最高，岷江冷杉的最低。岷江冷杉晚材宽度与年轮总宽度的相关性从1970年以后明显下降，而其他种的相关系数则随时间变化较小。树种之间标准化年表显著正相关，而云杉与紫果云杉和红杉与岷江冷杉之间相关系数明显较高。年表序列的第1主分量表达了4个树种树木共同径向生长变化格局；第2至第4主分量分别表达了云杉属和冷杉属、常绿针叶树种和落叶针叶树种以及云杉和紫果云杉树木径向生长变化差异。 不同海拔高度的8个岷江冷杉样地年轮宽度年表序列敏感度大体上随海拔高度升高而降低。各样地早材宽度与年轮总宽度年表之间的相关系数均在0.9以上，且随海拔高度变化不大；晚材宽度与年轮总宽度之间的相关系数随海拔高度的变化较大，并有随海拔升高而降低的趋势。样地年表序列之间相关系数差异很大，高海拔样地年表间多为显著正相关；低海拔样地年表间的相关系数变化不一；高海拔和低海拔样地年表之间相关性较差，且多不显著。样地年表的第1主分量能解释年表序列总方差的37.5%，反映了不同海拔高度岷江冷杉林木共同的径向生长变化格局；第2和第3主分量分别解释总方差的24.5%和18.2%，表现出明显的高海拔和低海拔样地树木间不同的径向生长变化，除一些样地例外，它们一般与低海拔样地年表有正相关，与高海拔样地年表有负相关。在那些另外的样地，海拔以外的其他因素可能也影响了树木径向生长变化。不同海拔高度样地林木的生长抑制和生长释放频率在不同时期表现出较大的差异，表明了不同的干扰历史和林木补充时间。 人工针叶林和次生混交林各样地林木早材宽度与其年轮总宽度年表之间相关系数均高达0.9以上；晚材宽度与年轮总宽度年表之间也都显著正相关，但人工针叶林样地的明显较高。样地年表序列之间的相关关系表现为，林分起源和经营管理相似的样地年表之间的相关系数明显较高，如人工针叶林与人工针叶林尽管树种不同，但样地年表之间显著正相关，而与次生混交林样地年表间关系不显著；反之亦然。综合比较各项生长参数及不同时期的树木径向生长速率，人工针叶林树木的胸径增长至少在40年以内是优于次生混交林的同种（或不同种）针叶树的。不同样地林木生长释放和生长抑制及人工针叶林树木胸高断面积增长分析表明，除严重的人为干扰外，林分郁闭后林木密度过大是造成高频率生长抑制的主要原因，在林分发育的适当时期通过抚育间伐等措施调控林分密度，是保证林木胸高断面积在一定时期内保持较高的连年增长的关键。日本落叶松作为引进的树种，在海拔3100 m左右种植表现良好，近30年来各项生长指标均高于林龄相近的云杉人工林，因此，适当用其作为川西亚高山针叶林采伐迹地快速恢复是合理的。 Miyaluo area is one of the typical regions covered by subalpine coniferous forests in western Sichuan province of southwestern China. To develop the regional tree-ring width chronology series for the dominant conifers such as Abies faxoniana, Picea asperata, P. purpurea and Larix potaninii, and to understand the radial growth patterns of conifers in Abies faxoniana natural forest stands at different altitudes, and in coniferous plantations and natural regenerated mixed stands in their different restoring processes as well, increment cores were sampled in the field together with conventional plots investigations for dendroecological analyses. After the increment cores being prepared according to standard procedures, the ring widths (total-ring and intra-ring widths) were measured with a WinDENDRO image-analysis system, and the measured tree-ring sequences were crossdated and quality-controlled with the software COFECHA. Using the software ARSTAN, we developed tree-ring width based chronology series of the four dominant conifers, eight site-specific Abies faxoniana chronologies, and seven site-specific chronologies of conifers in coniferous plantations and natural regenerated mixed stands. Mean sensitivities for total ring width chronologies of the four sampled dominant conifers were all below 0.2, while those for the latewood width chronologies of the same species were relatively much higher. Correlation coefficients between standard earlywood and total ring width chronologies of the four conifers were all above 0.9, but those between standard latewood and total ring width chronologies exhibited differences among species, with the coefficient of Larix potaninii the highest and that of Abies faxoniana the lowest. Correlation coefficients between latewood and total ring width of A. faxoniana obviously decreased from 1920-1970 for successive 50-year segments with 10-years lag analyses, though the same for the other three species changed unnoticeably with time. Tree-ring standard chronologies among species showed significant positive correlations, with the correlation coefficients between chronologies of Picea asperata and P. purpurea, and of Larix potaninii and Abies faxoniana relatively much higher. The first principal component of tree-ring chronologies represented the common radial growth patterns of the four conifers in Miyaluo area. The second, third and fourth PCs expressed the differences in radial growth responses for the genus Picea and Abies, for the evergreen and deciduous confers, and for the two species of the genus Picea, respectively. In general, mean sensitivities of the eight Abies faxoniana site-specific tree-ring width chronologies decreased with increasing altitude. The correlation coefficients between earlywood and total ring width chronologies for all sites reached 0.9, which did not change much with altitude; but those between latewood and total ring width chronologies diversified, with a decreasing tendency from lower altitudinal sites to higher altitudinal sites. Correlation coefficients among site chronologies varied considerably, with significant positive correlations among higher site chronologies, mixed results among lower site chronologies, and poor and insignificant correlations between chronologies of higher site and lower site. The first PC, which represents 37.5% of the total variance, reflected a common radial growth response at sites of different altitudes, and it showed a tendency of explaining more variance with increasing altitude. The second and the third PCs contributed to 24.5% and 18.2% of the total variance, respectively, exhibiting distinctive differences in radial growth responses between low- and high-altitudinal sites. With some exceptions, the radial growth represented by the second and third PCs had a positive correlation with that at the low-altitudinal sites and a negative correlation with that at the high-altitudinal sites. For those exceptional sites, factors other than altitude might also play a role in affecting tree-ring growth variations. Trees in stands of different altitudes showed great differences in frequencies of growth suppressions and releases through times, suggesting different disturbance histories and periods when trees recruiting to the canopy. Correlation coefficients between earlywood and total ring width chronologies for all sites of coniferous plantations and natural regenerated mixed stands were also above 0.9; and the same between latewood width and total ring width chronologies all positively correlated, too, with the coefficients of the coniferous plantations obviously much higher. Correlations among site chronologies showed that the coefficients among sites with similar stand origin and management regimes were much higher than those among sites with different stand origin and management regimes. For example, significant positive correlations were found for chronologies among different coniferous plantations, irrespective of species differences; while insignificant correlations between chronologies of the same conifer from a coniferous plantation and a natural regenerated mixed stand, and vise versa. Integrative comparisons of different tree growth parameters and radial growth rates at different stages indicated that the diameter at breast height (DBH) increments for trees in coniferous plantations were faster than those for trees of the same (or different) species in the natural regenerated mixed stands, at least within their first 40 years of stand development. Analyses of growth releases and suppressions, and basal area increments of trees in different stands demonstrated that over-dense individuals after canopy closure was the main factor resulting in high frequencies of radial growth suppressions, with some exceptions of severe man-made disturbances. Therefore, to ensure a continuous basal area current annual increment in certain periods, tree density controlling through thinning in due time during the stand development process are necessary. It should be mentioned that, as an introduced conifer to Miyaluo area, Larix kaempferi grew quite well at altitude of ca. 3100 m after planting in 1970s. In their near 30 years of stand development, Larix kaempferi trees exhibited faster growth in various parameters than Picea asperata trees of the similar stand age did. Thus we think it reasonable to use Larix kaempferi as a fast restoring species at appropriate sites of cutting blanks of subalpine coniferous forests in western Sichuan.

卧龙自然保护区不同海拔的中国沙棘(Hippophae rhamnoides subsp. sinensis)天然群体的遗传多样性分析

中国沙棘是一种雌雄异株、风媒传粉的灌木或乔木，在中国西南的卧龙自然保护区有广泛的分布。本研究以采集于四川卧龙自然保护区5 个海拔(1800 m、2200 m、2600 m、3000 m、3400 m)梯度的中国沙棘天然群体为材料，以ISSR 和AFLP 标记技术研究其遗传多样性水平及其遗传结构，旨在了解卧龙地区中国沙棘天然群体的遗传多样性水平以及遗传多样性在群体间、群体内以及雌雄亚群体间的分布和特征，为中国沙棘树种的遗传改良及种质资源保存提供遗传研究背景与实验依据。同时探讨ISSR、AFLP 和RAPD三种标记对中国沙棘天然群体的遗传变异水平和群体间遗传结构的评估能力和各自的优缺点。研究得出以下主要结论： 1. ISSR和AFLP分析都表明卧龙自然保护区的中国沙棘群体拥有较高的遗传变异水平(h = 0.249，HT = 0.305)。出现这种结果的主要原因可能与卧龙自然保护区多变的气候条件和生境的异质度大有关。 2. ISSR 和AFLP 都揭示出卧龙自然保护区中国沙棘群体的遗传多样性随着海拔的增加发生显著的变化，表现为中海拔群体(2200 m 和2600 m)比高海拔群体(3000 m 和3400 m)和低海拔群体(1800 m)有更高的遗传多样性的趋势。出现这种趋势的可能解释是低海拔群体处在相对高温和相对干旱的环境，高海拔群体受到低温和紫外线胁迫，而中海拔群体存在中国沙棘生长的适宜环境。 3. ISSR 和AFLP 分析都表明：卧龙自然保护区中国沙棘的遗传结构遵循分布范围广、交配系统以异交为主的木本植物的通常模式，即大多数的遗传变异存在于群体内，只有少部分的遗传变异存在于群体间。 4. 经Mantel 检测表明，卧龙自然保护区中国沙棘群体间的海拔距离和对应遗传距离之间存在显著的正相关关系，即随着垂直海拔距离的增加，群体间的遗传距离也随之增加。Mantel 检测结果以及聚类分析将卧龙自然保护区5 个不同海拔的中国沙棘群体分为低、中、高海拔群体三组的研究结果都表明，海拔很可能是限制群体间基因交流的主要因素。 5. ISSR 分析发现同一海拔的雌雄亚群体首先聚类的研究结果表明，同一海拔的雌雄亚群体在遗传上最相似。方差分析结果表明只有3.8%的总遗传变异存在于雌雄亚群体间，这可能与雌雄植株间的交配和遗传物质的混合有关。 6. ISSR、AFLP 和RAPD 分析都表明卧龙自然保护区不同海拔的中国沙棘天然群体的遗传多样性水平较高。它们的分析结果估算得到的Nei's 平均基因多样度(h)分别为0.249、0.214 和0.170。从该结果可以看出ISSR 和AFLP 比RAPD 检测到更多的遗传多态性,这很可能是不同标记检测的基因组的位点不同所致。 7. 依据对不同标记系统的比较分析，认为ISSR、AFLP 和RAPD 三种分子标记系统都能成功地用于调查卧龙自然保护区不同海拔的中国沙棘群体的遗传变异水平及遗传变异结构，提供关于中国沙棘天然群体多态性水平和遗传变异分布的有用信息。在三者中，AFLP 具有最高效能指数和标记指数，在确定种间分类关系或鉴别个体方面是一种比较理想的标记。 Hippophae rhamnoides subsp. sinensis, a dioecious and deciduous shrub species,occupies a wide range of habitats in the Wolong Nature Reserve, Southwest China. Ourpresent study investigated the pattern of genetic variation and differentiation among fivenatural populations of H. rhamnoides subsp. sinensis, occurring along an altitudinal gradientthat varied from 1,800 to 3,400 m above sea level in the Wolong Natural Reserve, by usingISSR and AFLP markers to guide its genetic improvement and germplasm conservation. And,comparative study of ISSR, AFLP and RAPD was performed to detect their capacity toestimating the level and pattern of genetic variation occurring among the five elevationpopulations of H. rhamnoides subsp. sinensis, and to discuss their application to the study onplant genetics. The results were list following: 1. The ISSR and AFLP analysis conducted for the H. rhamnoides subsp. sinensispopulations located in the Wolong Natural Reserve of China revealed the presence of highlevels of genetic variation (h = 0.249, HT = 0.305). Besides such features as relatively widedistribution, dominantly outcrossing mating system, and effective seed dispersal by small animals and birds, it is sometimes argued that hard climatic conditions and heterogeneous habitats may also contribute to high levels of diversity. 2. Genetic diversity of H. rhamnoides subsp. sinensis populations was found to varysignificantly with changing elevation, showing a trend that mid-elevation populations (2,200m and 2,600 m) were genetically more diverse than both low-elevation (1,800 m) andhigh-elevation populations (3,000 m and 3,400 m). H. rhamnoides subsp. sinensis is thoughtto be stressed by drought and high temperature at low elevations, and by low temperature athigh elevations. The high genetic variability present in the mid-elevation populations of H.rhamnoides subsp. sinensis is assumed to be related to a greater plant density in the middlealtitudinal zone, where favorable ecological conditions permit its continuous distributioncovering the zone from 2,200 m to 2,600 m above sea level. 3. The genetic structure of H. rhamnoides subsp. sinensis revealed by ISSRs andAFLPs followed the general pattern detected in woody species with widespread distributionsand outcrossing mating systems. Such plants possess more genetic diversity withinpopulations and less variation among populations than species with other combinations oftraits. 4. In the present study, Mantel tests showed positive correlations between altitudinaldistances and genetic distances among populations or subpopulations. The observedrelationship between altitude and genetic distances, and the result of the cluster analysisincluding populations or male subpopulations and classifying the groups into three altitudeclusters suggest that altitude is a major factor that restricts gene flow between populationsand subpopulations. 5. The analysis of molecular variance showed that only 3.8% of the variability residedbetween female and male subpopulations. Such a very restricted proportion of the totalmolecular variance between female and male subpopulations is due to common sexuality andmixing of genetic material between females and males. 6. The analysis based on ISSRs, AFLPs and RAPDs all revealed relatively high levelsof genetic variation among different altitudinal populations of H. rhamnoides subsp. sinensisin Wolong Natural Reserve of China. Their estimates of mean Nei’s gene diversity is equal to0.249, 0.214 and 0.170 respectively, suggesting the higher capacity of detecting geneticvariation of ISSR and AFLP than RAPD. It might be ascribed to their distinct sensitivity todifferent type of genetic variation. 7. Based on the coparative study on ISSR, AFLP and RAPD, we drew a conclusion thatthey all successfully reveal some useful information concerning the level and pattern ofgenetic vatiation occurring among different elevation populations of H. rhamnoides subsp.sinensis. AFLP is a ideal tool to taxonomic study and individual identification for theirhighest efficiency index and marker index among the three marker systems.