疏叶骆驼刺的生物生态学研究

盐生草甸是新疆平原草地生物生产力最高的植被类型。而豆科植物疏叶骆驼刺则是盐毕草甸植被的常见种类。此项工作对疏叶骆驼刺天然分布的范围及其生态环境条件，特别是土壤水盐状况和与该植物相联系的植物群落类型进行了调查分析。本文通过对K、Na、Ca、Mg4种金属盐性元素在土壤—疏叶骆驼刺系统中的迁移进行分析，探讨该种植物在盐化土壤环境中的生长状况与主要盐性金属元素的相互关系;通过对包括土壤植物体内元素含量、土壤水盐状况、植株生长状况等指标的综合分析，研究其生态适应关系和种群与土壤水盐状况的关系;通过对植物器官形态解剖学的研究，探讨该种植物与盐生土壤环境生态适应的生物结构基础和耐盐的生理途径;通过对土壤盐化过程影响因素和土壤盐化趋势的分析，结合疏叶骆驼刺的生态学特性，探讨该种植物在新疆盐化土壤的可能利用价值。结果如下： 1、疏叶骆驼刺为中生适弱盐化的潜水草甸植物。在有潜水发育的地带，其生存适应土壤盐化环境的范围较广。 2、在天然条件下，疏叶骆驼刺以根萌生新芽繁殖方式为主。在弱盐化和土壤水分较充足的地带，可以种子繁殖再生新株。 3、作为盐生草甸植被与荒漠植被间过渡类型中常出现的种类，疏叶骆驼刺的种群消长代表了一定的土壤盐化的演替阶段。 4、该种植物的器官组织显微结构显示出其与盐化土壤环境关系上特殊的适应机制。 5、在植物的生长发育过程中，金属盐性元素的迁移积累变化的性质表明，在一定条件下，该种植物对土壤盐性具有改善作用。 6、面对新疆盐化土地使用状况和土壤盐化退化的趋势，疏叶骆驼刺的生态利用潜势很大。

毛乌素沙地两种固沙禾草种子萌发和幼苗生长与环境的关系

干旱区和半干旱区生长的植物具有复杂的生存机制，以确保其能够在特定的环境中生存和发展。植物在干旱的荒漠条件下的生存，与其特殊的种子萌发机制密切相关，这种机制能够确保植物在合适的时间与地点进行种子萌发与幼苗生长发育。在植物的生活史中，种子对极端环境具有最大的忍耐力，而萌发的幼苗对环境胁迫的忍耐程度最小。在干旱区生长的植物往往具有特殊的萌发机制使萌发出的幼苗能够度过对外界的敏感期，对于植物的生存具有重要意义。 毛乌素沙地是我国的四大沙地之一，该地区具有水分短缺，蒸发强烈，风沙剧烈和生境异质性高的特征。本文假设生长在这种极端环境中的植物也发展出了“适时适地”的种子萌发和幼苗生长的适应对策。为了验证以上的假设，本文选取毛乌素沙地不同生境中生长的两种优势固沙禾草——流动沙丘上生长的沙鞭(Psammochloa villosa)和固定沙丘上生长的赖草(Leymus secalinus)为研究材料，通过野外调查、温室控制实验和实验室控制实验的方法，从生理生态学的角度探讨这两种植物的种子萌发和幼苗生长过程对沙丘环境的适应对策，主要对比它们在种子休眠、萌发和幼苗早期生长过程中对沙丘生境适应性的异同点。研究结果表明： （1）新成熟的沙鞭和赖草的种子为适应冬季低温而发展出生理性的内生休眠——非深度生理休眠。沙鞭和赖草的种子分别需要经过4周和8周的低温层积处理(3－5ºC)来完全打破休眠。另外，划破种皮或者部分切除胚乳也能够促进种子的萌发，这进一步证明两种植物的种子具有非深度生理休眠。然而，切除胚乳在不同程度上影响它们的幼苗生长。由非深度生理休眠、温度和损伤种皮/胚乳调节的部分萌发机制能够确保两种植物的种子即使在条件适宜的情况下只有部分种子萌发，从而分散植物生存的风险性。 （2）毛乌素沙地的小量降水（无法触发萌发）使种子经常遭受湿润－干燥的交替胁迫过程。种子先在湿润条件下吸涨1d或者2d，然后在室温下干燥0-8天。尽管在经历反复吸涨和自然干燥脱水后仍能够保持萌发能力，沙鞭和赖草种子的萌发特性却发生了不同的变化：和各自的对照相比，沙鞭种子萌发率相同而萌发速率降低;赖草种子的萌发率和萌发速率都降低，部分种子进入休眠状态。沙鞭和赖草萌发出的幼苗可能由于没有后续降雨或者因沙蚀而遭受干燥胁迫，但是其幼苗在生长早期能够忍耐一定程度的干燥，再次湿润后部分幼苗能够恢复生长。沙鞭和赖草幼苗的耐干燥的“极限点”不同：当幼根长度为1 mm时，它们的幼苗忍耐干燥的时间分别是60d和30d;当幼根长度为4 mm时，它们的幼苗忍耐干燥的时间分别是14d和7d。沙鞭和赖草的种子和生长早期的幼苗的耐干燥性特性可能是它们对降雨量和降雨时间都不可预测的沙地生境的生存策略之一。 （3）不同的沙埋深度影响沙鞭和赖草的种子萌发和出苗。这两种植物的种子萌发和出苗都需1－2 cm的浅层沙埋。随着沙埋深度的增加，两种植物的种子萌发率和出苗率逐渐降低，强迫休眠率逐渐升高;萌发率与出苗率和沙埋深度呈负相关关系而休眠率和沙埋深度呈正相关关系。但是，沙鞭种子出苗的最大沙埋深度是8 cm，而赖草的则为4 cm。因强迫休眠而没有萌发的种子对维持一个长期的土壤种子库来说具有生态学优势，这些种子暴露在合适的萌发土壤深度时具有生长出幼苗的潜能。 （4）沙鞭和赖草的种子都具有大小的差异性，种子大小对沙鞭和赖草的种子在不同沙埋深度的出苗具有不同的影响。沙鞭的三种不同大小种子的平均质量分别为小，4.489 ± 0.012 mg (4 – 4.9 mg);中，5.457 ± 0.012 mg (5 – 5.9 mg)和大，6.415 ± 0.011 mg (6 – 6.9 mg)。赖草的两种不同大小种子的平均质量分别为小，3.083 ± 0.026 mg (3 – 3.5 mg)和大3.955 ± 0.028 mg (3.6 – 4.0 mg)。在相同的沙埋深度下，两种植物的大种子的出苗率都显著高于小种子。和小种子相比，两种植物的大种子由于贮藏更多的能量，所以在相同深度的沙埋中具有出苗率更高的生态优势，而大量小种子在沙埋中不能萌发，可以作为种子库保存在沙层中，这样就分散了一次性大量萌发给植物带来的冒险性。 （5）沙鞭和赖草的幼苗在生长过程中会遭受沙埋，其幼苗忍耐沙埋的能力与沙埋的相对深度（沙埋比例）和幼苗年龄有关。沙鞭和赖草幼苗的耐沙埋能力不同：沙鞭的2周龄幼苗可以忍耐达到株高100%的沙埋，而其1周龄幼苗只能忍耐75%的沙埋。赖草的1周龄和2周龄幼苗都只能忍耐75%的沙埋。沙埋之后，沙鞭和赖草幼苗的生物量，根/茎比以及根和茎的长度都受到不同程度的影响。赖草幼苗不能忍耐完全沙埋可能是限制它在流动沙丘上分布的一个原因。 （6）降雨量和降雨频率能够不同程度地影响沙鞭和赖草在不同沙层的萌发和出苗。这两种植物的种子萌发和出苗需要的最小降雨量不同：在一次浇水相当于5 mm降雨量后，沙鞭和赖草种子的萌发率都超过50%;但是使沙鞭和赖草的出苗率能够达到50%的降雨量分别为10 mm和15 mm。沙埋中的沙鞭和赖草种子的出苗对降雨的响应具有以下特征：两种植物种子的出苗随降雨量或者降雨频率的增加而增加;沙鞭的出苗率受到降雨量和降雨频率的显著影响，但是二者交互作用的影响不显著;赖草的出苗率受到降雨量、降雨频率以及二者交互作用的显著影响。 由非深度生理休眠，种子大小，干燥－湿润循环，沙埋和降雨调节的种子萌发和出苗机制确保了自然条件下沙鞭和赖草每次只有少量种子萌发和出苗，从而分散了两种植物在沙丘上的生存风险。 根据沙鞭和赖草在沙丘上的种子萌发和幼苗生长特性，本文为毛乌素沙地通过植物固沙恢复受损的沙地生态系统的种子飞播实践提出了几点建议。

神农架巴山冷杉种子雨及种实特性研究

巴山冷杉主要分布于秦巴山地，是湖北神农架地区暗针叶林的建群种，构成了当地的优势群落，对神农架地区的植被景观具有决定意义。研究巴山冷杉的结实特性，种子雨时空格局，种子萌发能力和活力劣变过程，有利于进一步认识和分析这一物种的生活史特征，群落组成结构和更新动态。对这一重要物种的合理保护和有效利用具有促进作用。 在湖北神农架自然保护区选择巴山冷杉－箭竹、巴山冷杉－茵芋和巴山冷杉－陕甘花楸群落设置样地，通过布设种子收集器结合室内实验分析，对巴山冷杉种子雨的时空格局进行研究。待巴山冷杉球果成熟时，人工采集球果和种子，测量和比较巴山冷杉的结实特征，利用发芽实验研究巴山冷杉的萌发特性，并通过人工加速老化实验探讨巴山冷杉种子的劣变过程。 巴山冷杉种子雨始于10月初，持续超过2个月，不同类型的巴山冷杉种群表现出不同的种子雨时空格局。巴山冷杉－箭竹群落中巴山冷杉种子雨平均强度167.93±111.14粒/m2，有活力种子比例占22.31%，落种高峰期集中于10月27日到11月2日间，种子雨在种群中呈现聚集分布。巴山冷杉－茵芋群落种子雨强度在三个样地中最小，只有16.41±14.41粒/m2，有活力种子仅占3.05%。巴山冷杉－陕甘花楸群落，种子雨落种高峰为10月15日至10月21日，林冠内种子雨数量占到了收集总数的87.95%。扩散的种子数量与离开中心母树的距离间的关系近似正态分布。 对巴山冷杉的结实特性的分析结果表明，巴山冷杉球果近圆柱形，球果平均长5.37±0.75cm;平均宽3.01±0.32cm;重量介于5.9g～41.3g。巴山冷杉种子长1.08～8.68mm，种子宽1.16～6.42mm，种子厚0.66～3.48mm，种子千粒重7.30g。球果单果出种量在不同种群的频数分布格局不同，巴山冷杉球果和种子的各项指标在种群间和种群内存在显著差异。球果大小、出种量及单粒种子的质量间存在着极显著的正相关关系。此外，种子生活力水平在母树间差异较大，不同母树的种子发芽率和发芽势不等且随时间的萌发进程也不相同，相同种源的巴山冷杉种子在不同的培养条件下萌发能力不同，室温条件下具有最高的发芽率但在25℃，8h光照，恒温培养时的萌发最为整齐。 高温高湿条件能够加速巴山冷杉种子的老化进程，使巴山冷杉种子的含水量发生变化，种子的活力水平随着老化时间的延长而降低，40℃，100% 相对湿度处理6天后种子基本丧失活力。

薇甘菊对其入侵地植物种子萌发和幼苗生长的化感影响

薇甘菊主要入侵我国华南的广东、海南和云南三省，给当地造成了严重的危害，其强烈的化感作用被认为是其成功入侵的原因之一。而目前对薇甘菊化感作用的研究还较少，因而对薇甘菊化感作用的认识还有待深入的研究。另外，在对入侵植物入侵群落进行恢复时，由于入侵植物的凋落物以及生长过的土壤都还含有化感物质，常常导致恢复难以继续。因而选择抗化感能力强的物种来恢复薇甘菊的入侵地也成为该研究的一个主题。本文以薇甘菊为实验材料来研究薇甘菊化感作用对植物种子活力、种子萌发和幼苗生长的影响，同时筛选化感抗性强的物种以进行薇甘菊入侵地的恢复工作。其主要结论如下： 1、薇甘菊根和叶的提取液对植物种子活力、种子萌发和幼苗生长都有很大的抑制作用，并且这种抑制作用随着提取液浓度的升高以及处理时间的延长而加强，这证明了薇甘菊具有很强的化感作用。 2、一些植物种子萌发和幼苗生长在低、中浓度提取液以及高浓度提取液的短时间的处理下能被促进，不过在高浓度的提取液以及长时间的处理下都被抑制，这说明化感作用是抑制还是促进作用在很大程度上受到提取液浓度和处理时间的影响。 3、薇甘菊提取液（50 g L-1）对紫薇(Lagerstroemia indica)、刺槐(Robinia pseudoacacia)、黑麦草(Lolium perenne)和西葫芦（Cucurbita pepo）种子处理50-100 h就能极大的影响到其后的种子萌发和幼苗生长，这说明薇甘菊在短时间（50-100 h）就能给植物产生极强的化感作用，也进一步证实薇甘菊很强的化感作用。 4、由于各物种进化历史的不同，而且各物种在对薇甘菊的化感抗性上也存在着差异，所以各物种对薇甘菊提取液的化感反应也各不相同。 5、薇甘菊根提取液对刺槐和紫薇幼苗根的生长的影响要大于叶提取液对刺槐和紫薇幼苗根的生长的影响，而对刺槐和紫薇种子萌发率、初始萌发时间、萌发速率以及幼苗株高的影响却是叶提取液的化感作用强。这支持了一个新的假设：由于物种间代谢的多样性，至少在根和叶对受试植物根的化感影响方面，对于一些物种来说根的化感作用可能强于叶的化感作用，对于另一些物种来说叶的化感作用可能强于根的化感作用。 6、薇甘菊叶提取液对西葫芦种子活力、种子萌发和幼苗生长的化感影响大，而对其后开花时间以及开花的数目没有明显的影响，这说明薇甘菊对西葫芦生活史前期的化感影响要大于对西葫芦生活史后期的化感影响。 7、薇甘菊叶的水提取液对种子萌发的抑制作用要大于对幼苗生长的抑制作用;对非豆科植物种子萌发的影响要大于对豆科植物种子萌发的影响，而对非豆科植物幼苗生长的影响要小于对豆科植物幼苗生长的影响;对本地种的抑制作用要大于对外来归化种的抑制作用。 本研究证实了薇甘菊强烈的化感作用，根提取液对受体植物幼根生长的影响大于叶提取液对受体植物幼根生长的影响，这种化感作用可能极大的促进了薇甘菊的入侵。另外，在恢复薇甘菊入侵群落时，考虑到薇甘菊的化感作用，理应选择外来的已经归化了的非豆科植物的幼苗。

毛乌素沙地四种关键植物幼苗出土及生长对沙埋和水分的响应

飞播是毛乌素沙地植被恢复与重建的重要手段。但此项技术仍存在一些问题，如飞播后成苗率较低等。柠条(Caragana korshinskii)、羊柴(Hedysarum laeve)、籽蒿(Artimisia sphaerocephala)与油蒿(Artimisia ordosica)是毛乌素沙地广泛分布的物种，也是飞播的主要物种。本文以飞播植被恢复技术为突破口，研究沙埋和土壤水分对种子萌发，幼苗出土和生长的影响，探讨幼苗出土和生长对沙生环境的适应对策。对采用合适的物种进行生态工程建设，提高生态工程的稳定性及可持续性具有重要意义。同时，本项研究有助于加深半干旱区植物对沙生环境适应机理的认识。 以上述4种植物为研究对象，设定7个沙埋深度和9个水分梯度，研究种子萌发和幼苗出土对沙埋和单次供水的响应。设定7个沙埋深度，选择3个沙丘部位，研究沙丘不同部位对植物种子萌发的影响。设定6个沙埋深度、4个水分梯度，研究两种蒿属植物幼苗生长对不同沙埋和供水的响应。得出如下主要结论： (1) 对于种子质量相对较大的柠条和羊柴而言，在一次性10-20 mm供水量条件下，幼苗主要在0.5-2 cm埋藏深度之间出土，但在埋藏深度为5 cm时这两个物种仍有少部分幼苗能够出土;对于种子质量相对较轻的油蒿和籽蒿而言，在一次性10-20 mm供水量时，幼苗主要在0.5 cm埋藏深度出土，油蒿和籽蒿幼苗在埋藏深度为1.5和2 cm时不能出苗。沙埋深度为0.5 cm时，4个物种出苗率较高、出苗时间较短、出苗速率较快。 (2) 实验期内(30天)在一次性供水量分别为5、7.5和7.5mm时，柠条、羊柴和油蒿出土幼苗全部死亡;在一次性供水量高于15-20 mm时，出土幼苗死亡率低于50%;籽蒿在供水量高于10 mm时出土幼苗死亡率低于50%;随供水量的增加4个物种出土幼苗的死亡率降低。 (3) 结合毛乌素沙地降雨特点与本文实验结果，0.5 cm的沙埋及7.5 mm以上的单次降雨量是上述4种植物自然条件下出苗较高的重要条件。 (4) 在沙丘顶部，2-7 cm沙埋深度之内各个物种不同沙埋深度的种子萌发率差异不显著。沙丘背风坡中部和底部，4个物种在沙埋深度为1 cm时的种子萌发率均显著高于5、7 cm时的萌发率。当沙埋深度为1-5 cm时，柠条和羊柴种子在坡面不同位置的萌发率均高于20%，且显著高于7 cm沙埋深度时的萌发率。在背风坡底部时，柠条、羊柴、油蒿和籽蒿的种子萌发率达到最大值，分别为45.2±3.27%、48.4±5.21%、20.8±4.63%和22.4±4.83%。4个物种的种子萌发率从沙丘顶部到背风坡中部到底部呈递增趋势，表明背风坡中部和底部的环境条件要比顶部更适合种子萌 发。 (5) 部分沙埋促进幼苗生长。2种蒿属植物幼苗在沙埋深度为0.25、0.5 H时比在0、0.75 H时具有更高的生物量、叶面积、相对生长速率（RGR）和净同化速率（NAR）。在75 mm/月供水量时，油蒿幼苗具有较高的生物量和叶面积，但当供水量达到100 mm/月时其幼苗生长受到抑制。油蒿幼苗在遭遇较深（0.5、0.75 H）沙埋时的根冠比要显著高于未沙埋幼苗的根冠比;籽蒿幼苗根冠比在不同沙埋条件下没有显著变化。籽蒿幼苗在50 mm供水量时具有较高的RGR。油蒿幼苗在75 mm供水量时具有较高的RGR和NAR。

濒危植物大花黄牡丹种子休眠与萌发特性研究

本文以中国极危种大花黄牡丹种子为研究材料，对其种子生物学、休眠与萌发特性进行研究，采用变温层积和激素处理解除种子休眠，并通过生物抑制物测试、胚培养、激素含量动态变化的测定，研究种子休眠的原因，从根本上解决了大花黄牡丹迁地保护过程中种子繁殖的技术难题，并初步探讨了大花黄牡丹致濒因素与种子休眠及萌发特性的关系，结果表明： 1.大花黄牡丹种子饱实度高，活力强，种皮较坚硬，但不存在吸水障碍，干燥条件下易因失水而丧失活力。迁地保护冷室中盆播18个月方可出苗，出苗率约4%;变温层积实验表明该种具有典型的下胚轴休眠和上胚轴生长抑制。 2.胚组织培养表明：带胚乳和种皮的胚不能萌发，剥除种皮和胚乳后，胚根可萌发，胚芽不生长，GA3处理上胚轴可促进生长，说明种皮和胚乳是导致大花黄牡丹种子下胚轴休眠的关键因素，而上胚轴生长抑制与胚本身关系更密切。 3.抑制物质提取实验表明：大花黄牡丹种子胚乳、种皮、胚等各部位的浸提液均存在抑制小白菜种子萌发的物质，且抑制作用依次增强，说明胚乳和胚是其下胚轴休眠的主要原因。经过暖层积（15 ℃/90 d）种子（未解除上胚轴生长抑制）的胚根、子叶、胚轴浸提液对小白菜和已解除休眠的大花黄牡丹种子的萌发均有不同程度的抑制作用，并依次减弱，说明种胚本身的抑制物质是导致生理休眠的主要原因。 4.变温层积处理和外源激素实验表明：新鲜种子采收后15 ℃暖层积3个月生根率可达85%，下胚轴休眠解除对温度要求严格，高于或低于15 ℃及变温条件均不利于下胚轴萌发;暖层积90d、根长大于6 cm种子再经过60～80 d/ 5 ℃冷层积，即可有效解除大花黄牡丹种子上胚轴的生长抑制，出芽率达80%，最终出苗率68%。不同浓度GA3及不同处理时间促进上胚轴伸长实验结果显示，GA3　400 mg/L浸种根长大于1.5 cm的种子2 h，出芽率可达100%，可以完全解除上胚轴的生长抑制作用。 5.休眠萌发过程中种子各部位内源激素含量动态变化分析结果说明，初始状态脱落酸含量高是导致大花黄牡丹种子下胚轴休眠、上胚轴生长抑制的主要原因之一，且上胚轴抑制与子叶、下胚轴和根的脱落酸含量密切相关;同时认为种子各部位初始生长素含量水平低是导致其休眠的另一个主要原因;变温层积过程中胚各部位脱落酸含量的急剧下降和生长素的迅速升高是解除休眠的关键;同时发现赤霉素在解除休眠和促进萌发过程中起着重要的促进作用，外源GA3能够有效地打破上胚轴的深度休眠。玉米素核苷在休眠与萌发进程中变化趋势与生长素和赤霉素相似，说明其对种子胚根和上胚轴的萌发和生长具有一定的促进作用。

Effects of dietary cyanobacteria of two different sources on growth and recovery of hybrid tilapia (Oreochromis niloticus x O. aureus)

A 115 days feeding trial was conducted to evaluate the effect of dietary cyanobacteria on growth, microcystins (MCs) accumulation in hybrid tilapia (Oreochromis niloticus x O. aureus) and the recovery when the fish were free of cyanobacteria. Three experimental diets were formulated: the control (cyanobacteria free diet); one test diet with cyanobacteria from Lake Taihu (AMt 80.0 mu g MCs g(-1) diet) and one with cyanobacteria from Lake Dianchi (AMd, 410.0 rho g MCs g(-1) diet). Each diet was fed to fish for 60 days and then all fish were free of cyanobacteria for another 55 days. A significant increase in feeding rate (FR) was observed in fish fed AMd diet after a first 30-day exposure (1(st) EP), and in fish fed both AMt diet and AMd diet after a second 30-day exposure (2(nd) EP). Specific growth rates (SGR) of fish fed AMt diet and AMd diet were both obviously affected after the first 30-day exposure, but SGR was only significantly affected in fish fed AMt diet after the second 30-day exposure. After a 55-day recovery, there were no significant differences among diets in the indices mentioned above. Much higher concentrations of MCs were accumulated in tissues of all fish exposed to cyanobacteria. After the 55-day recovery, MC concentrations in fish tissues were significantly lower than those on day 60. (C) 2009 Elsevier Ltd. All rights reserved.

黄土丘陵植被恢复过程中土壤种子库萌发数量特征及动态变化

采用野外定点采样与室内土壤种子萌发试验相结合的研究方法,选取黄土丘陵区典型草原带宁夏固原地区不同植被恢复年限(2,5,12,24,35,55,75 a)及当年农地(对照)的8个样地,研究该区土壤种子库萌发数量特征及动态变化。结果表明,8个不同植被恢复年限样地土壤中可萌发的种子的平均数量为3542.5粒/m2,可萌发种子以双子叶植物为主,约占萌发总数的90%。0~5 cm土层可萌发种子数量变化于166.7~9100.0粒/m2;5~10 cm土层变化于133.38~2120.02粒/m2。土壤中可萌发种子数量随着恢复年限的增加表现先增加后下降的趋势。土壤中可萌发的种子数量随时间的变化近似单峰型,双子叶植物峰值主要出现在第4~6周,而单子叶植物萌发高峰在5~6周。表明在环境,气候等条件适宜时,黄土丘陵区典型草原带可萌发的土壤中的种子可形成萌发高峰。

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。

水分亏缺和凋落物水浸液对云杉幼苗生长和种子萌发的影响

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

川西亚高山云杉人工林天然更新与生境异质性研究

川西亚高山针叶林是四川森林的主体，是长江上游重要的生态屏障。云杉作为亚高山针叶林人工更新的主要树种，已经在该地区形成了大面积的人工纯林。目前，许多云杉人工林分已经进入更新成熟龄，而这些人工林的持续更新却成为日益凸现的问题。探讨这些云杉人工林的自我更新潜力及云杉种子种群更新特点，可为培育后续森林资源提供科学依据。 本文以川西米亚罗亚高山60a云杉人工林为研究对象，并以该区域内相对稳定的植被群落——天然林为对照，采用种子收集器、土壤种子库筛选、室内外种子萌发实验及野外幼苗调查等方法，从异质性微生境的角度研究了种子雨下落之后，不同微生境对种子库、种子萌发、幼苗建立及分布这一前期更新过程的影响，得出如下结果： 1、通过对川西亚高山60a云杉人工林和天然林6年内种子雨雨量、形态特征、散步动态等的持续观测和综合比较可以发现，云杉林结实特点由于林木自身的特征存在着巨大的变动，2002年和2006年两个种子结实大年内，60a人工林种子雨强度分别达到1088.2 ± 52.3粒/m2和704.3 ± 48.9粒/m2，远大于天然林579.9 ± 28.9粒/m2 和507.5± 30.7粒/m2；且云杉林结实质量优于天然林。60a人工林结实量大，种子质量也最好，相对天然林来说对种群的天然更新以及群落的演替都有最大的贡献潜力。应该说，在川西亚高山云杉人工林的天然更新过程中，种源不是影响天然更新的因素。在种子结实大年里，达到更新成熟的云杉人工林有着优于该地区相对稳定植被群落——天然林的种源优势。至少在种子结实大年，种子供应不是该区域人工林天然更新的限制因子。 2、相对于天然林种子库，人工林种子库在种子萌发前能够有较多的有活力种子。虽然这其中有种子雨输入量有差别的因素存在，但两种林分种子库对种子的保存率的不同才是造成这种差异的主要因素。在人工林中，不同地被类型形成的微生境显著地影响了种子库中种子的密度、垂直分布。有地被物存在的微生境能够将种子雨的大部分截留在地被层中，成为幼苗出现的主要场所；同时不同的地被物对种子的保存情况存在显著的差异，苔藓和凋落物层能都较好地保持其中的种子，到种子萌发前，这两种种子库类型能分别为天然更新提供366.1粒/m2和302粒/m2的有效种子。从这点来看，林下地被物上的种子库能够为天然更新萌发阶段提供数量可观的物质基础。 3、种子的萌发和幼苗的定居是天然更新过程中种子库向幼苗库转化的关键环节。总的说来，米亚罗人工林区60a云杉人工林种子向幼苗的转化率十分低下，凋落物、苔藓、草本、裸地四种主要地被物以及天然林内种子/幼苗的转化率分别仅为2.22%、2.14%、0.57%、0.67%、1.05%。这种低的转化率成为云杉林天然更新的限制性因子。但在现有更新条件下，微生境对这一环节仍然显示出十分显著的影响，表现为凋落物和苔藓对现有更新的新幼苗的保存率高于其它类型及天然林。苔藓和凋落物在种子萌发，幼苗保存，以及幼苗分布上都要优于其它地被物类型；另外，微地形对天然更新过程的影响也很显著，凹立地上更适宜于种子的汇集、萌发和定居。 Subalpine coniferous forests dominate most parts of the forested areas in western Sichuan, and they are important ecological barriers in the upper reaches of the Yangtze River. Picea asperata is one of the keystone spruce species for re-afforestation after felling of the natural forests and there have been a total of ca. 13 000 ha of plantations dominate by this species established. Nowadays, many P. asperata plantations have reached reproductive maturity. However, continued regeneration becomes to an important problem in these plantations. Understanding their self-regeneration potential and the regeneration characteristics of seed populations in spruce plantations of these plantations can have some insights on the management of these plantations and the establishment of following forest resources. A subalpine 60a P. asperata plantation distributed in Miyaluo artificial forest area was studied in this paper, at the same time. Synchronously, a 150a natural spruce forest was studied as comparison. Using seed collecting traps, sieving method for soil seed bank, seed germination experiments and seedling investigations in the field, the effects of heterogeneous microsites on early natural regeneration processes after seed rain were studied, which included seed banks, seed germination, seedling establishment and distribution. The main results are as follows. 1. Through a 6-year long term investigation of seed rain intensities, characteristic, dispersal dynamics of 60a P. asperata plantation, we could concluded that the seed setting properties of 60a P. asperata plantation have a great variation for the characteristics trees. In the mast seed year of 2002 and 2006, the seed rain intensities of plantation was 1088.2 ± 52.3 seeds/m2 and 704.3 ± 48.9 seeds/m2 respectively, which were much greater than that of natural spruce forest (579.9 ± 28.9 seeds/m2 in 2002, and 507.5± 30.7 seeds/m2 in 2006). Furthermore, the quality of seed rain in P. asperata plantation was better than that of natural spruce forest. Contrasting with natural spruce forest, 60a P. asperata plantation has a greater potential on natural regeneration of P. asperata population and succession of community for the reason of greater seed rain intensities and better seed quality. We can confirm that seed source was not a limiting factor which influences the natural regeneration progress of P. asperata population distributed in subalpine mountain zone, at least in the mast seed year. 2. Contrasting with natural spruce forest, P. asperata population had more viable seeds in seed bank before the germination. Although the difference of seed rain intensities of two forests has effect on this phenomenon, the difference of seed conservation ability in two forests was the main factor. In the P. asperata plantation, the seed densities and seed vertical distribution pattern were significant effected by the microsites, which posed by different ground cover types. In other word, Microsite with ground covers can obstruct most seeds and keep them in ground cover layer from seed rain, and these ground covers would be the main site for seed occurrence. There was a significant difference about seeds conservation ability among these ground covers. Litter and moss could better conserve P. asperata seeds which distributed in this two covers. Seed banks exist in litter and moss ground cover types could respectively provide 302seed/m2 and 366.1seed/m2 for natural regeneration before the seed germination. From this point of view, we could conclude that ground covers can ensure considerable numbers of seeds for the germination process. 3. Seed germination and seed establishment are key steps that the seeds invert to seedlings in natural generation process. In sum, the seed/seeding transform rate in 60a P. asperata plantation distributed in Miyaluo artificial forest area is very low. the seed/seeding transform rates in litter, moss, herb, soil surface and natural spruce forest were 2.22%、2.14%、0.57%、0.67%、1.05%, respectively. The low transform rate become to a limiting factor of P. asperata natural regeneration process. However, under the existing conditions of natural regeneration, microsit still had significant effect on this transform. The states of Seed germination, new seedling conservation, and older seedling distribution in litter and moss were better than in any other ground cover type or natural spruce forest. In addition, the micro-relief has significant effect on natural regeneration process. Concave site was more suitable for seed collection, seed germination and seedling distribution.

花椒水浸液及酚酸类化感物质对作物种子萌发和幼苗生长的影响效应

花椒（Zanthoxylum piperitum）是川西干旱河谷地区的重要经济作物，化感作用是花椒连作障碍的原因之一。系统研究花椒化感作用有助于深入理解并最终解决花椒连作障碍。本文通过研究花椒叶、林下土壤浸提液及单一纯化感物质对花椒幼苗生长、苜蓿种子萌发及幼苗生长的影响作用，从生理生化角度揭示浸提液及纯化感物质的作用方式。通过室外和室内模拟实验，对浸提液及纯化感物质的化感效应进行比较，为花椒连作障碍的解决和化感作用机制的深入理解提供依据。主要结果如下： 1.花椒叶及林下土壤浸提液对地下生物量影响作用强与对地上生物量的化感效应，两种浸提液的化感效应强度不同，叶浸提液作用表现更显著。其中在Y6、Y8 、T6和T8处理时，花椒幼苗地下生物量分别降低了31.2%、32.1%、31.6%和31.7%。 2.两种浸提液均能显著影响花椒幼苗体内的保护性酶活性，总体说来，在高浓度下抑制各种抗氧化物酶活性，幼苗体内丙二醛含量增加，幼苗受害严重；在较低浓度下，各种保护性酶活性有所增加，丙二醛含量减少，幼苗伤害减轻。同时，不同月份里，各种酶的活性高低显著不同，9月份的活性显著低于7月份的酶活性。对于养分含量的影响，Y8、T8的影响强度最大，分别使碳元素含量降低了27.8%和30.8%，使钾元素含量降低了34.7%和25.6%。 3.花椒叶及林下土壤浸提液对苜蓿种子萌发及幼苗生长有化感作用，表现在最终萌发率、不同物质代谢及保护性酶活性的差异上。两种浸提液对苜蓿种子萌发过程中蛋白质的含量均无显著性影响，对淀粉和可溶性糖的影响作用类似，高浓度处理无明显化感效应，较低浓度处理显著降低二者在萌发苜蓿种子中的含量。Y2、Y4与T4处理分别使可溶性糖含量减少了32.3%、29.1%和18.8%，Y2与T2处理分别使淀粉含量降低了29.3%和26.8%。 4.苜蓿种子在4种单一化感物质最高浓度即10-3 mol•L-1处理下，萌发率显著降低，半数萌发时间推迟，随着处理浓度降低，抑制作用逐渐减弱，当降低到10-6 mol•L-1时，又能够表现出对苜蓿种子萌发的促进作用。 5.纯化感物质在10-6 mol•L-1时使苜蓿幼苗叶片的保护性酶活性显著升高，丙二醛含量显著降低；在10-3 mol•L-1时使苜蓿叶片中保护性酶活性显著降低，丙二醛含量增加，膜脂过氧化程度加重。 Zanthoxylum piperitum is one of the most important cash crops and has been extensively cultivated in Eastern Tibetan Plateau, especially in the fragile dry valley areas. Allelopathic effects could be a reason for Z. piperitum’s continuous cropping impediment. Systemmatically research of the effect of Z. piperitum allelopathy could help to comprehend the continuous cropping impediment. The allelopathic effects on seedlings growth and seed germination of aqueous extracts of Zanthoxylum piperitum and phenolic allelochemicals were studied, and the action mechanism of the two substances was also discussed from physiology. Indoor and outdoor experiments were set to compare the difference between aqueous extracts and pure allelochemicals. The main results showed that: 1. The aqueous extracts of leaf and soil had significant allelopathic effects on aboveground and underground biomass, but the effect on underground biomass was stronger than the effect on underground evidently. Treated with Y6、Y8 、T6 and T8, the underground biomass was reduced 31.2%、32.1%、31.6% and 31.7% respectively. 2. The activity of activities of superoxide dismutase, catalase, peroxidase and ascorbate peroxidase were significantly reduced, while the content of MDA was increased and the seedlings were suffered stronger, when treated by the high concentration; but at the low concentration, these were reversed. And then, at the different month, the activities of antioxidant enzyme were significantly distinct. As for the contents of nutrient element, Y8、T8 had the more intensive effects than other treatments. 3. The results showed that the two types of aqueous extracts had significant allelopathic effects on seed germination, substances metabolize and the activity of antioxidant enzyme. But the aqueous extracts had no effects on the content of protein, while had the similar effects on the content of starch and soluble sugar. At Y2、Y4 and T4, the content of soluble sugar decreased 32.3%、29.1% and 18.8% respectively. 4. Treated with 10-3 mol•L-1 of the four allelochemicals, the seed germination of alfalfa was significantly inhibited. Ferulic acid, coumarin and vanillic acid at 10-3 mol•L-1 significantly reduced the activities of antioxidant enzyme, while the content of MDA in alfalfa seedling was significantly increased. The restrain effects became weakened with the treat concentration falled. However, ferulic acid, coumarin and vanillic acid could increase the activities of antioxidant enzyme at 10-6mol•L-1.

岷江上游高山林线附近优势树种土壤种子库和幼苗更新特征的研究

岷江上游地区高山/亚高山植被分布的坡向性分异显著，阴阳坡高山林线不仅物种组成差异明显，并且分布海拔呈现出阴坡高阳坡低的格局．阳坡林线树种主要是圆柏属乔木，林线类型多为渐变型，海拔高度大约在3 400m~3 800m；阴坡林线树种主要是冷杉，林线类型多为骤变型，海拔高度约在3 800m~4 400m．本研究采用土壤种子库物理筛选、室内萌发实验及野外群落调查等方法，对岷江上游地区阴坡岷江冷杉和阳坡祁连圆柏两类林线树种不同海拔梯度上土壤种子库以及幼苗库特征进行了调查，从土壤种子库和幼苗更新特征的角度对林线乔木树种种群更新特征进行了分析，进而对该地区高山林线在阴阳坡分布差异的原因进行了探讨，结果显示： 1．土壤种子库 阴坡：阴坡高山林线附近岷江冷杉土壤种子的平均密度大约为50.96粒/m2，其中树线以上10m处土壤种子密度为1.00粒/m2，树线处大约19.33粒/m2，林线交错带内土壤种子密度最高为136.83粒/m2，郁闭林内种子密度小于林线交错带，只有30.50粒/m2，种子平均空壳率为52％，霉变率达34%，完好种子只有6%．土壤种子库垂直分布特征为地被物层含种子比重最大，大约在67.50%左右；其次为0~2cm层，约18.84%左右；2~5cm层所占种子比例最小，约13.66%左右．霉变种子数量与土壤深度呈负相关． 阳坡：阳坡祁连圆柏土壤种子的平均密度为60.16粒/m2．树线以上10m处密度为1.92粒/m2，树线位置大约108.16粒/m2，林线交错带内平均为75.80粒/m2，郁闭林内种子密度小于林线交错带，只有20.00粒/m2．种子平均空壳率为36％，完好种子占49%，霉变率较低，大约为10%．阴阳坡林线树种土壤种子库垂直分布特征为：地被物层含种子最多，其次为0~4cm层，4~10cm层所占种子比例最小，霉变种子数量与土壤深度也呈负相关． 2． 幼苗库调查 阳坡：在树线以上区域没有发现幼苗，林线交错带内幼苗密度平均达3 250株/hm2，郁闭林内仅2 750株/ hm2．整个样地内1～2a幼苗很少甚至没有出现，3～10a的幼苗相对较多．空间分布上，祁连圆柏幼苗在林线交错带内接近随机分布，郁闭林内则介于随机分布和均匀分布之间． 阴坡：在树线以上幼苗密度为1 250株/ hm2，全部为1~2a幼苗，林线交错带内幼苗密度平均达7 000株/ hm2，郁闭林内达6 250株/ hm2．林线附近岷江冷杉幼苗丰富度以及幼苗的出现频率明显高于祁连圆柏，年龄结构也较祁连圆柏完整．岷江冷杉幼苗空间分布除了树线处幼苗的分布为随机分布，其他海拔则为集群分布． 3．从不同土壤深度的种子总量和幼苗数量的相关性检验发现，当年生幼苗数量跟表层种子总量相关性极显著, 但是两年生幼苗的数量与底层种子数量相关性显著．土壤种子在土壤中的垂直分布格局从一定程度上可以反映种子库的年际特征．岷江冷杉土壤种子库较丰富，种子散布后的存活力随着时间的变化逐渐下降，属于季节性瞬时种子库；祁连圆柏土壤种子散布格局为集群型分布，成熟种子大部分散布在母株冠幅内，属于永久性土壤种子库． 4．在阴坡林线交错带及以上区域还存在较为丰富的乔木土壤种子，并且在树线以上区域还发现了少量的岷江冷杉幼苗．从样地乔木的年龄结构发现，在林线交错带内上部到树线位置主要以幼龄林为主，且年龄结构完整，基本符合入侵性林线特征；阳坡林线交错带内幼苗出现频率很低，树线以上区域虽然存在种子库，但是没有幼苗出现，在林线交错带内乔木径级差距很大，年龄结构异常不完整，这种特征的林线将会面临两个可能结果：一种是维持现有状态，保持平衡；另外一种就是退化，但阳坡林线的实际动态趋势还有待长期定点研究． Treelines on the upper region of Minjiang River differ between the north aspect and the south aspect in their appearances, altitudes and tree species. On the north aspect, trees of Abies form a sharp and abrupt treeline ranging from 3800m to 4400m, while on the south the treeline is generally lower(3 400~3 800m), more open and gradual and mostly composed of Sabina. In this study, we examined the altitudinal gradients of soil seed banks and seedling recruitments at the treeline ecotones of a N-aspect and a S-aspect by using soil sieving, germination experiment and field investigations, analyzed the characteristics of population regeneration of tree species at the transitional zone and presented a analysis of the causes to the aspect-related difference in treeline patterns in the study area. Major results of our study include: 1． Soil seed bank N-aspect: Of the 50 plots investigated, the average density of soil seeds is 50.96/m2, in which well-formed seeds account for 6%, empty seeds 52%, parasitized seeds34%, and seeds damaged by animals 8%. The size of soil seed bank varies along altitude, being 1.00 seeds /m2 at the 10m above the treeline and ca.19.33 seeds/m2 at the upper limit of treeline. The highest density (136.83 seeds/m2) occurs at the treeline ecotone. By contrast, the density of soil seed for the closed forest is only 30.50 seeds/m2. In terms of vertical strata, 67.50% of the total seeds are at the surface layer, 18.84% at the middle layer (0~2cm) and 13.66% at deeper layer (2~5cm). The number of parasitized seeds is negatively correlated to soil depth. S-aspect: Of the 50 plots investigated, the average density of soil seeds is 60.16 seeds/m2, and the well-formed seeds account for 49%, empty seeds 36%, parasitized seeds10%, and seeds damaged by animals 1%. The size of soil seed bank varies along altitude, with 1.92 seeds/m2 recorded at the10m above the treeline，108.16 seeds/m2 at the upper limit of treeline, and 75.80 seeds/m2 at the treeline ecotone, while that for the closed forest is 20.00 seeds/m2. The number of seeds decreases with the depth of soil. As is on the N-aspect, the size of soil bank, from large to small, follows the order of the surface layer, the middle layer (0~4cm) and the bottom layer (4~10cm). The number of parasitized seeds is also negatively correlated to the depth of the soil. 2． Seedling bank N-aspect: A mean maximum seedling abundance of 31 000 seedlings/hm2 was recorded near alpine treeline at growing season. The density of seedlings is 1 250 seedlings/ha (all being 1 or 2 years old) at the alpine meadow 10m away above treeline, 7 000 seedlings/ha at treeline ecotone and 6 250 seedlings/ha for closed forest．The spatial distribution of Abies faxoniana seedlings is random at the upper limit of the treeline but clumped at other altitudes． S-aspect: No seedlings were found at the alpine meadow 10m away from the treeline. The density of seedlings was 3 250 seedlings/ha at treeline ecotone and 2 750 seedlings/ha for the closed forest．Hardly any 1 year current and 2 year-old seedlings appeared at the plots. The spatial distribution of Sabina przewalskii seedlings is random at treeline ecotone and between “random” and “even” forest closed forest． 3．Correlation tests of seedling population and seed bank at different soil layers indicated that the emergents were strongly correlated to seed bank at surface layer while the number of two-year seedlings was significantly correlated to the seed bank at the bottom of soil layer, indicating that germination mainly occurs at the soil surface while the middle or bottom layer was the reserve for non-germination or dead seeds. It can thus be postulated that Abies faxoniana soil seed bank is of seasonal transient type. By contrast, the soil seed bank of Sabina przewalskii is of persistent type and the soil seeds and seedlings of this species occurred more frequently near the islands of adult trees. 4．A good many soil seeds of both tree species were found near the treeline ecotone and above at N- and S-aspects. A few young seedlings were found above the Abies treeline. Investigation of five altitudinal transects respectively on N- and S-aspects indicated that Abies faxoniana has a more complete age structure than the stands of Sabina przewalskii. The age of firs decreased from closed forest to the upper limit of treeline, which suggests that the Abies treeline is advancing to higher altitude. While on the south aspect, only few Sabina przewalskii soil seeds and nearly no seedlings were found above the treeline ecotone. The stands exhibit extremely great difference in diameter classes with significantly incomplete age structure. This would lead to two possible results for the treelines: maintaining an equilibrium state at the current position or degenerating. But more studies should be carried out at longer time scales or larger spatial scales to understand whether the Sabina treeline is degenerating.

化感物质香豆素对苜蓿种子萌发及幼苗初级氮同化的影响

香豆素类物质是苯丙酸内酯（环酯）类化合物，绝大部分高等植物通过次生代谢途径都能合成。研究表明，香豆素类物质是花椒体内最重要的化感物质，系统研究香豆素类物质的作用机理有助于理解和最终解决花椒连作障碍。本文通过研究香豆素对几种植物种子特别是苜蓿种子萌发、苜蓿幼苗初级氮同化的影响，从生理生化角度揭示香豆素的作用方式，为花椒连作障碍的解决和化感作用机制的深入理解提供依据。主要研究结果如下：1. 研究了香豆素对6 种常见作物种子萌发的影响，并对一组数据采用4个不同的指标进行评价，对生物测定化感作用中存在的问题进行了讨论。结果发现1.0mM的香豆素对采用的6 种作物的种子萌发均表现出一定的化感作用，4 个指标的敏感程度依次为S (发芽速度)＞AS(累积发芽速度)＞CRG(发芽指数)＞GT(最终发芽率)。种子萌发实验是化感作用研究中最重要、应用最广泛的生物测定方法之一，应根据不同的研究目的合理采用指标和实验方法。2. 采用培养皿试纸法进行种子萌发试验，研究了香豆素水溶液在苜蓿种子萌发过程中对其吸水、电导率及抗氧化保护酶活性的影响。结果表明，影响苜蓿种子发芽的香豆素浓度阀值为0.3mM。香豆素在1.0mM 的浓度下降低了苜蓿种子吸水阶段Ⅱ的吸水速度，使其外渗物质增多，电导率增大，并显著抑制了超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)的活性，同时种子体内丙二醛（MDA）的含量显著增大。高浓度香豆素破坏了膜的结构、影响了抗氧化保护酶的活性是香豆素降低苜蓿发芽率的原因之一，也可能是影响花椒-苜蓿间作的关键因素之一。3. 不同浓度（0、25 μM、50 μM、0.1 mM、1.0 mM）化感活性物质香豆素对10 日龄苜蓿幼苗初级氮同化的影响的结果表明25 µM～50 µM 的香豆素加快了苜蓿幼苗对硝态氮的吸收。高浓度的香豆素导致苜蓿根系和叶片内可溶性蛋白含量降低、鲜重减小、地下鲜重/地上鲜重（R/S）的比值升高，根系中初级氮同化的关键酶硝酸还原酶（NR）、谷氨酸胺合成酶（GS）、谷氨酸脱氢酶（GDH）的活性降低，叶片中NR、GS 的活性减低、叶绿素含量减少，而GDH 的活性升高。香豆素影响苜蓿幼苗氮代谢和氨同化的关键酶，导致体内养分的缺失是香豆素抑制苜蓿幼苗生长的机理之一。Coumarins are lactones of o-hydroxycinnamic acid, and are allelopathiccompounds that originate in the phenylpropanoid pathway. They are synthesized byalmost all higher plants. According to previous studies, coumarins were mostimportant allelochemicals in Chinese prickly ash. Systematically research of theeffect of coumarin could help to comprehend the continuous cropping impediment.The effects of coumarin on seed germination and primary nitrogen assimilation ofalfalfa were studied. The main results showed that:1. We compared four common germination indices (S, AS, CRG, GT)preciously calculated with the same date. The results showed that, at theconcentration of 1.0 mM, coumarin inhibited seeds germination. Among all indices,the S index was most sensitive, followed by the AS and CRG indices. Andsuggestions on the expression of bioassay results were also provided.2. At concentrations above 0.3 mM, coumarin inhibited seed germination in aconcentration-dependent manner. During seed imbibitionⅡ, coumarin at 1.0 mMsignificantly reduced the activities of superoxide dismutase (SOD), catalase (CAT),peroxidase (POD), while the content of malonyldialdehyde (MDA) in alfalfa seedssignificantly increased. The higher concentration coumarin destroyed structure ofmembrane and influenced activities of antioxidant enzymes, which might be one ofthe reasons that coumarin decreased germination rate of alfalfa, and one of the keyfactors influencing Chinese prickly ash-alfalfa intercropping.3. Alfalfa plants were exposed to different concentration of coumarin (0、25μM、50 μM、0.1 mM、1.0 mM) grown for 10 days on control medium. Coumarin, in the range of 25 μM～50 μM, significantly stimulated the net nitrate uptake.Increasing coumarin concentration led to a decrease of protein contents in theleaves and roots. The root to shoot (R/S) FW ratio was increased by increasingcoumarin concentration. Under high coumarin concentration, the activities of nitratereductase (NR) and glutamine synthetase (GS) were repressed in the roots andleaves. Glutamate dehydrogenase (GDH) was inhibited in the roots, while enhancedin the leaves. Chlorophyll contents in the leaves were also decreased under highcoumain concentration. Coumarin decreased alfalfa growth by (i) nutritionaldeficiencies shown by the decrease of nitrate, (ii) lowered N compound synthesisvia inhibition of nitrate reduction and ammonium assimilation.

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

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