论我国兴安落叶松、长白落叶松及华北落叶松种间遗传进化关系

摘要/Abstract

摘要：

兴安落叶松、长白落叶松及华北落叶松是我国重要的木材树种,然而由于采用不同的性状进行分类,它们种间关系问题一直在争论着。国外研究人员如Ostenfeld and Larsen(1930)等认为长白落叶松和华北落叶松为兴安落叶松的2个变种,而我国科学家则把它们视为两个单独的种,并且还认为长白落叶松有3个变种存在,而华北落叶松有1个变种存在。因此用分子遗传学方法研究兴安落叶松、长白落叶松及华北落叶松3个种间遗传进化关系有着重要的实际意义,例如种间杂种优势的利用潜势预测。至今植物分类研究所采用的性状已包括有表型形态、生殖、生化、蛋白质(同功酶)到RNA和DNA的分子标记等,进行综合性状分类比较。由于这些变量所经受的环境影响和所包含的各种历史事件(如突变、选择、连锁等)不同,因此应用不同性状分类时,有可能导致不一致的结果。本文分别从形态性状,包括球果和木材解剖特征等、染色体核型分析、同功酶变异、RAPD分析、叶绿体DNA的RFLP分析,以及叶绿体DNA3个非编码碱基序列(1452bp)对这3个落叶松种的遗传进化关系进行了详细的探讨和评述。从表型和生态适应性状变异看,这3个落叶松种可以较容易的区别出来,它们有着不同的生态特性和若干形态学差异,然而这可能只反映了适应性性状(受自然选择影响)的变异。近年来应用同功酶标记(选择性中性标记)研究结果表明:这3个种的遗传关系非常密切,种间Nei氏遗传距离平均约为0.01,种内群体间的遗传距离约为0.002。3个落叶松的种间遗传距离要比其它落叶松种间距离要小,如用同功酶分析所得的欧洲落叶松(L.decidua)和西伯利亚落叶松(L.sibirica)Nei氏距离为0.057。应用叶绿体DNA分子标记研究结果表明3个落叶松种间无差异。这意味着它们的遗传分化可能发生很晚、或种群间的基因流动发生十分频繁。根据所用的同功酶变异特点,即选择性中性和检测后证明为连锁平衡(Hu,1998),若分化时间尺度很短的话,那么突变的影响可以忽略不计,这时遗传漂变和基因流动对群体的遗传变异起主导作用。研究结果显示平均单位点的等位基因数和多态性水平均由兴安落叶松、长白落叶松到华北落叶松依次减少,由此可推测长白落叶松与华北落叶松可能的形成过程。然而这一推测的前提是突变影响很小,而这一前提可以从叶绿体DNA的变异以及LePage和Basinger根据古化古资料对其它落叶松种的分化推测间接加以判断。根据分子钟(Molecular Clock)假设和Juke与Cantor-参数模型,可以粗略地估算出这3落叶松的分化时间发生很晚,可能发生在第三纪后期(Late Tertiary),这要比叶绿体DNA发生突变频率的倒数要小,这样也就间接地推断突变的影响很小。虽然对于生物的概念有许多不同的观点,如基于分子标记而建立的系统进化种与传统的生物学种可能不一致,然而我们根据生物学上种的概念以及一些性状变异特点,认为将长白落叶松和华北落叶松划为兴安落叶松的2个变种而非2个单独的种可能要更合理些。这与Shiraishi et al.(1996)建议将发生在Mt.Manokami山上的落叶松划为日本落叶松(L.kaempferi)的一个变种而非一新种的情况类似。根据这种非常近的遗传关系,我们认为3个落叶松种间的杂种优势利用是非常有限的。为了进一步研究3个落叶松种的形成与遗传关系,我们建议应用线粒体DNA标记进行更深入的研究。虽然线粒体DNA的点突变率要比叶绿体DNA要低,但由于线粒体DNA分子内存在高频率的重排(Rearrangement),进而产生遗传变异。同时也需要了解到这种变异不适合于分析种间的进化关系(Phylogeny Reconstruction),及线粒体DNA的母本遗传方式特点,但可以用群体遗传结构的研究。稀有标记的空间分布有助于进一步判断落叶松群体的形成路径和历史。

关键词: 兴安落叶松, 长白落叶松, 华北落叶松, 遗传进化关系

Abstract:

The three taxa of the genus,Larix gmelinii(Rupr.)Rupr,L.olgensis Henry and L.principis-rupprechtii Mayr.,are very important timber species in China. However,the classification of these three taxa has been very arguable because different traits are employed by different authors. Thus,elucidation of their genetic relationships is of great singnificance in practice,such as the potential utility of heterosis between the taxa. Recent studies assessed using allozymes and cpDNA markers showed that there were very closely genetic relationships among the three taxa,implying that their divergences probably occurred very recently. In this review the probable process for the formation of L.olgensis and L.principis-rupprechtii was discussed in detail,including possible routes,the time scale for divergence and the definition of species. According to the concept of biological species and the genetic variation of different traits,it would be more reasonable to consider L.olgensis and L.principis-rupprechtii to be two varieties of L.gmelinii other than two separate species.

Key words: L.gmelinii(Rupr.)Rupr, L.olgensis Henry, L.principis-rupprechtii Mayr., Genetic evolutionary relationship

胡新生,Ennos Richard A.,王笑山. 论我国兴安落叶松、长白落叶松及华北落叶松种间遗传进化关系[J]. 林业科学, 1999, 35(3): 84-96.

XIN-SHENG HU,RICHARD A. ENNOS,XIAO-SHAN WANG. ON EVOLUTIONARY RELATIONSHIPS AMONG THREE LARIX TAXA IN CHINA:L.GMELINII(RUPR.)RUPR, L.OLGENSIS HENRY AND L.PRINCIPIS-RUPPRECHTII MAYR.[J]. Scientia Silvae Sinicae, 1999, 35(3): 84-96.

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