马尾松和黄山松物种分化遗传机制研究
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摘要
物种分化遗传机制是当前生物学研究领域的一个热门话题。由于生殖隔离在物种分化过程中起着十分重要作用，阐明物种间生殖隔离的遗传机制是物种分化研究领域的一个重要目标。然而迄今物种生殖隔离产生的原因仅在少数几个物种中进行了较为深入的研究。种间杂交往往会在杂种的育性及适应性方面产生负面效应，即使一些物种通过种间杂交能产生杂种优势，但在后续世代的杂交中往往也会发生不育现象。遗传作图为我们揭示亲本基因组中控制配子向后代不均等传递的基因组区域提供了重要的技术手段。在遗传作图过程中，配子向后代不均等传递表现为标记偏分离。由于配子不均等传递的显著程度与物种分化程度存在正相关关系，因此在种间杂交组合中，检测标记偏分离的基因组区域可用于查找控制种间生殖隔离有关的基因组区域。目前这类研究主要是利用人工杂交组合展开，利用天然杂种进行的研究还鲜有报道。
马尾松与黄山松的物种分化主要是由于它们所处的海拔生态位不同造成的，它们在垂直分布上相临，且存在自然的基因交流，因此马尾松与黄山松是研究临域性物种分化遗传机制的理想材料。
在本文中，作者系统研究了马尾松、黄山松及其杂种个体的球果特征、种子的饱满度、发芽情况一些生理生化指标的变化规律。另外，由于松属基因组硕大，笔者针对松属树种进行了AFLP分析程序的优化。在此基础上，利用AFLP标记和一株马尾松与黄山松杂种个体上采集的种子的胚乳组织构建了杂种个体的遗传图谱。利用建成的遗传图谱，分析了可能与这两个树种生殖隔离相关的基因组区域，并进一步利用数量遗传学手段，定位了控制标记偏分离的遗传位点。最后利用表达序列标签将建成的连锁图连结到松属树种的共祖先图谱上。上述研究为加深我们在基因组水平上对物种分化遗传机制的认识提供了重要的实验证据。论文开展的上述几个主要方面的研究内容及结果如下：
1．不同海拔松树球果形态及种子质量指标分析：以采自安徽黄山风景区不同海拔的松树球果、种子为研究材料，测定了球果长、球果宽、球果长宽比、种子饱满度、发芽率和千粒重6项指标，并分析了这些指标随海拔高度的变化情况。结果表明：随着海拔的升高，球果长、球果宽、长宽比减小；杂种域个体种子的饱满度、发芽率、千粒重3个指标的均值小于黄山松和马尾松。在所有采集球果的植株中，种子饱满度随海拔高度呈现两头高，中间低的变化趋势。其中海拔800米以下个体种子平均饱满度约为73%，海拔1100米以上个体种子平均饱满度为57.6%，而800-1100米之间的个体种子平均饱满度只有40%。过渡带种子的饱满度低（空粒率高）可能与马尾松和黄山松存在种间繁殖障碍有关。这一结果显示前人对马尾松、黄山松及其杂种域海拔分布的划分比较吻合。
2．黄山松幼苗生理生化指标随海拔高度的变异情况：以不同海拔高度采集的黄山松种子所培养的幼苗为研究对象，将幼苗置于0-5℃冰箱中进行寒冷处理24h后，测定了幼苗的相对电导率、SOD、丙二醛、POD以及可溶性蛋白5个生理生化指标处理前后的变化情况。采用多重复测定，探讨了这些指标随海拔高度的变化关系。结果表明，幼苗浸出液相对电导率、SOD活性及丙二醛质量摩尔浓度在低温处理下，均表现出随海拔高度的增加而降低的变化趋势。这三个指标与黄山松的海拔适应性存在相关，这一工作为研究马尾松与黄山松分化遗传机制候选基因的选择提供了实验依据。
3．松属树种AFLP实验程序优化及引物组合筛选：AFLP标记技术可利用较少的DNA获取大量的位点分离数据，是胚乳组织进行遗传作图的理想标记技术。但松属树种的基因组硕大，超出了一般AFLP分析程序适用上限的10倍左右，因此在松属树种中利用AFLP标记技术，需要进行针对性的实验程序优化。这部分工作针对松属树种进行了AFLP分析过程中DNA酶切、接头连接、预扩增及选择性扩增实验程序优化。实验结果发现, E+4/M+3的引物组合比较适合于松属树种的AFLP分析。在这部分工作中，笔者筛选出了适合松属树种遗传分析的AFLP引物组合，为利用松属树种胚乳组织构建遗传图谱提供了标记资源。
4．马尾松、黄山松杂种遗传图谱构建及标记偏分离分析：遗传标记在杂交子代中的不均等传递是研究杂交不亲合遗传机制的重要手段。通过多年多地点采样，笔者发现这两个树种杂种域个体种子发芽率普遍偏低，反映两个树种存在一定程度上的生殖隔离。笔者利用一株马尾松与黄山松杂种个体上采集的种子得到的192份胚乳组织进行杂种个体的遗传图谱构建。标记偏分离分析发现，显著偏分离的标记比例极高，约占分离标记的25.5%。笔者进一步对标记偏分离的影响范围，标记偏分离的方向及标记偏分离的显著程度进行了分析，这些指标在不同连锁群上差异很大。这部分的研究结果揭示马尾松与黄山松的分化并不是全基因组水平上的分化造成的，而是局限于某些染色体上的局部区域。建成的图谱揭示了相关的基因组区域，并为相关遗传因子的基因组学解析提供了重要的信息整合平台。
5．偏分离遗传位点定位及相关基因组区域确定：由于配子不均等传递的显著程度与物种分化程度存在显著的相关关系，因此在种间杂交组合中检测标记偏分离的基因组区域可用于查找与控制种间生殖隔离有关的基因组区域。但以往受偏分离座位影响的基因组区域都是通过偏分离标记的数量及偏分离程度进行大致的估测。要确定目标基因组区间，需要对偏分离座位进行遗传定位分析。在这一部分工作中，笔者利用建成的马尾松与黄山松杂种遗传图谱，进行了偏分离位点定位分析，并将建成的遗传图联结到松属的共祖先图谱上。分析中共发现了6个LOD支持度强的SDL位点。这些位点分布于4个不同的连锁群上。由于这些位点控制作图亲本产生的配子向后代的不均等传递，SDL对应的基因组区域比基因组其它区域在黄山松与马尾松物种分化过程中应起着更为重要的作用。SDL分析是在远缘杂交组合中解析控制配子不均等传递遗传因子重要环节，这部分工作为SDL分析提供了一个研究实例，也是广大演化生物学家普遍感兴趣的一个研究内容。
Speciation is one of the most interesting processes in evolution, and reproductive isolationis essential for the process of speciation to be complete. Thus, understanding the geneticmechanism triggering reproductive isolation is a key goal in the study of speciation. To date, theunderlying causes of reproductive isolation are only partially understood in a few species.Hybridization between different species often has negative consequences that result in reducedfertility or viability, which leads to transmission ratio distortion (TRD) of the parental gametes.Occasionally, hybridization can raise hybrid vigor in which a hybrid offspring performs well.Yet, recombination and allelic segregation may render hybrid breakdown in the next generation.Genetic mapping methods provide a unique opportunity to study the interactions ofdifferentiated genes and genomes in a hybrid genetic background. The degree of TRD isthought to be positively correlated with the level of genome divergence between taxa, mappingthe SDLs that control the TRD marker clusters can be used to infer the genome regions relatingto reproductive barriers in distant crosses. To date, the majority evidences of TRD occurring inhybrid genetic backgrounds are based on studies employing experimental hybrids. By contrast,map-based studies of natural hybrids are much rarer.
Pinus hwangshanensis and P. massoniana are native conifers in south of China. In theareas to the south of Yangzi River, P hwangshanensis is mainly found in the elevation range of800-1800m, and P. massoniana distributes below elevation of800-900m. The major differencein the ecological niches of the two species is elevation. With an increase in elevation,environmental factors, such as oxygen partial pressure, air temperature and moisture regime,soil temperature and water regime, sunray and ultraviolet light intensity, will change. Theseenvironmental factors are closely related to plant growth and fitness. They are environmentalstresses to cause differentiation in plant phenology and fitness, subsequently, to maintain thespecies-specific characteristics of the alternate species. The distribution ranges of P.hwangshanensis and P. massoniana are frequently found to be adjacent to each other, andoverlapped with a narrow contact zone. This distribution pattern leads them to be the desirableresearch materials to uncover the genetic mechanism triggering the parapatric speciationprocess. In my thesis, studies were carried out in the following aspects: Variation in conemorphology and seed characteristics for P.massoniana, P. Hwangshanensis, and their hybridzone; Variation trends of physiological and biochemical indexes of P. hwangshanensis withincrease in elevation; AFLP protocol optimization and primer combination screening for Pinusspp.; Potential chromosomal introgression barriers revealed by linkage analysis in a hybrid of P. massoniana and P. hwangshanensis; Genome-wide detection of genetic loci triggering unevendescending of gametes from a natural hybrid pine. The main context include:
1. Variation in cone morphology and seed characteristics for P. massoniana, P.hwangshanensis and their hybrid zone. The cones and seeds were collected from differentelevations of Mt. Huang in Anhui Province. Six characteristics of cones and seeds, includingcone length, cone width, ratio of cone length to width, seed plumpness, germinationratio and weight per thousand seeds were analyzed to reveal the variation trends with thechanging of elevations. The results indicated that: Cone length, cone width, ratio of conelength to width decreased with increase in elevation; Seed plumpness, germination ratioand weight per thousand seeds were lower for plants in hybrid zone than that of theother zones. We propose it may relate to the reproduction barrier between P. massonianaand P. hwangshanensis. It was noticeable that the majority of variation for cones andseeds arose among individual trees, and traits were commonly diverged amongindividuals at different elevations.
2. Variation trends of physiological and biochemical indexes of P. hwangshanensiswith increase in elevation. P. hwangshanensis seeds collected from different elevations ofHuanggang Mountain in Wuyi Nature Reserve were cultivated in a container at25℃. Aftersprouting，the seedlings were stored in a refrigerator within a temperature range of2-5℃for24hours. Five physiological and biochemical indices of P. hwangshanensis seedlings weredetermined before and after the treatment，and the variation trends of these indexes withincreasing elevation were also studied. Three of the tested indices，including relative electricalconductivity of extract fluid，SOD activity，and MDA content of the seedlings after lowtemperature treatment decreased with increasing elevation. Therefore，the above three indicesare related to the adaptability of P. hwangshanensis to elevation. These indices could be appliedto explore the mechanism of parapatric speciation.
3. AFLP protocol optimization and primer combinations screening for Pinus spp.Pines possess gigantic genomes (10-20Gb), which are about10fold that of the human genomeand about40fold that of the poplar genome, leading them out of the genome range suitable forthe common AFLP(Amplified Fragment Length Polymorphism) protocol. This paper speciallyoptimized an economic and reliable AFLP analysis protocol for pines. The number of selectivenucleotides added to the core primers and the primer combinations suitable for AFLP analysisin such species were tested. Some AFLP primer combinations that were screened their utilityacross different species were tested. The results could also be used as a reference for allowingbetter amplification in complex templates isolated from plants species with large genomes
4. Potential chromosomal introgression barriers revealed by linkage analysis in ahybrid of P. massoniana and P. hwangshanensis. Distortion of marker transmission ratio isfrequently ascribed to selection against alleles that cause hybrid incompatibility. The naturalintrogression between P. massoniana and P. hwangshanensis and their distribution ranges lead to the emergence of the two species as desirable organisms to study the genetic mechanisms forspeciation. Using seeds sampled from trees at different elevations, we consistently detectedsharp decreases in seed germination rates of trees in the hybrid zone, which might be duelargely to the hybrid incompatibility. A genetic map was established using192megagametophytes from a single tree in the hybrid zone of the two species. Segregationdistortion analysis revealed that the percentage of significant-segregation-distortion (SSD)markers was extremely high, accounting for more than25%of the segregating markers. Theextension range, the distortion direction, and the distortion intensity of SSD markers also varieddramatically on different linkage groups. This study displays the potential chromosomalintrogression barriers between P. massoniana and P. hwangshanensis. The established mapprovides a valuable platform for conducting genome-wide association of hybrid incompatibleQTLs and/or candidate genes with marker transmission ratio distortion in the hybrid.
5. Genome-wide detection of genetic loci triggering uneven descending of gametesfrom a natural hybrid pine. Marker transmission ratio distortion (TRD) revealed in geneticmapping studies of distant crosses can be used to infer the genetic basis relating to reproductivebarriers between species. Unlike measure the degree of TRD by the overall number ofsegregation distorted markers in the affected genome regions, mapping the segregationdistorting loci (SDL) provides reliable statistic parameters that help to confine the targetgenomic region for further characterization at molecular level. Using the linkage mapconstructed for a natural hybrid of P. hwangshanensis and P. massoniana, we performed SDLanalyses and aligned the established map to the pine consensus map. Altogether,6SDLs withrelatively strong LOD supports were detected on4linkage groups of the established map. Sincegametes inheriting different alternate chromatid blocks from the SDL affecting genome regionshad uneven chance to descend to the offsprings, the corresponding genome regions weresupposed to play more significant role in rendering the speciation of P. hwangshanensis and P.massoniana. This paper presents a case study on a crucial step for uncovering the geneticfactors that triggered the uneven descending of gametes in a natural distant cross, and it maydraw broad interest for the evolutionary biologists.
马尾松与黄山松的物种分化主要是由于它们所处的海拔生态位不同造成的，它们在垂直分布上相临，且存在自然的基因交流，因此马尾松与黄山松是研究临域性物种分化遗传机制的理想材料。
在本文中，作者系统研究了马尾松、黄山松及其杂种个体的球果特征、种子的饱满度、发芽情况一些生理生化指标的变化规律。另外，由于松属基因组硕大，笔者针对松属树种进行了AFLP分析程序的优化。在此基础上，利用AFLP标记和一株马尾松与黄山松杂种个体上采集的种子的胚乳组织构建了杂种个体的遗传图谱。利用建成的遗传图谱，分析了可能与这两个树种生殖隔离相关的基因组区域，并进一步利用数量遗传学手段，定位了控制标记偏分离的遗传位点。最后利用表达序列标签将建成的连锁图连结到松属树种的共祖先图谱上。上述研究为加深我们在基因组水平上对物种分化遗传机制的认识提供了重要的实验证据。论文开展的上述几个主要方面的研究内容及结果如下：
1．不同海拔松树球果形态及种子质量指标分析：以采自安徽黄山风景区不同海拔的松树球果、种子为研究材料，测定了球果长、球果宽、球果长宽比、种子饱满度、发芽率和千粒重6项指标，并分析了这些指标随海拔高度的变化情况。结果表明：随着海拔的升高，球果长、球果宽、长宽比减小；杂种域个体种子的饱满度、发芽率、千粒重3个指标的均值小于黄山松和马尾松。在所有采集球果的植株中，种子饱满度随海拔高度呈现两头高，中间低的变化趋势。其中海拔800米以下个体种子平均饱满度约为73%，海拔1100米以上个体种子平均饱满度为57.6%，而800-1100米之间的个体种子平均饱满度只有40%。过渡带种子的饱满度低（空粒率高）可能与马尾松和黄山松存在种间繁殖障碍有关。这一结果显示前人对马尾松、黄山松及其杂种域海拔分布的划分比较吻合。
2．黄山松幼苗生理生化指标随海拔高度的变异情况：以不同海拔高度采集的黄山松种子所培养的幼苗为研究对象，将幼苗置于0-5℃冰箱中进行寒冷处理24h后，测定了幼苗的相对电导率、SOD、丙二醛、POD以及可溶性蛋白5个生理生化指标处理前后的变化情况。采用多重复测定，探讨了这些指标随海拔高度的变化关系。结果表明，幼苗浸出液相对电导率、SOD活性及丙二醛质量摩尔浓度在低温处理下，均表现出随海拔高度的增加而降低的变化趋势。这三个指标与黄山松的海拔适应性存在相关，这一工作为研究马尾松与黄山松分化遗传机制候选基因的选择提供了实验依据。
3．松属树种AFLP实验程序优化及引物组合筛选：AFLP标记技术可利用较少的DNA获取大量的位点分离数据，是胚乳组织进行遗传作图的理想标记技术。但松属树种的基因组硕大，超出了一般AFLP分析程序适用上限的10倍左右，因此在松属树种中利用AFLP标记技术，需要进行针对性的实验程序优化。这部分工作针对松属树种进行了AFLP分析过程中DNA酶切、接头连接、预扩增及选择性扩增实验程序优化。实验结果发现, E+4/M+3的引物组合比较适合于松属树种的AFLP分析。在这部分工作中，笔者筛选出了适合松属树种遗传分析的AFLP引物组合，为利用松属树种胚乳组织构建遗传图谱提供了标记资源。
4．马尾松、黄山松杂种遗传图谱构建及标记偏分离分析：遗传标记在杂交子代中的不均等传递是研究杂交不亲合遗传机制的重要手段。通过多年多地点采样，笔者发现这两个树种杂种域个体种子发芽率普遍偏低，反映两个树种存在一定程度上的生殖隔离。笔者利用一株马尾松与黄山松杂种个体上采集的种子得到的192份胚乳组织进行杂种个体的遗传图谱构建。标记偏分离分析发现，显著偏分离的标记比例极高，约占分离标记的25.5%。笔者进一步对标记偏分离的影响范围，标记偏分离的方向及标记偏分离的显著程度进行了分析，这些指标在不同连锁群上差异很大。这部分的研究结果揭示马尾松与黄山松的分化并不是全基因组水平上的分化造成的，而是局限于某些染色体上的局部区域。建成的图谱揭示了相关的基因组区域，并为相关遗传因子的基因组学解析提供了重要的信息整合平台。
5．偏分离遗传位点定位及相关基因组区域确定：由于配子不均等传递的显著程度与物种分化程度存在显著的相关关系，因此在种间杂交组合中检测标记偏分离的基因组区域可用于查找与控制种间生殖隔离有关的基因组区域。但以往受偏分离座位影响的基因组区域都是通过偏分离标记的数量及偏分离程度进行大致的估测。要确定目标基因组区间，需要对偏分离座位进行遗传定位分析。在这一部分工作中，笔者利用建成的马尾松与黄山松杂种遗传图谱，进行了偏分离位点定位分析，并将建成的遗传图联结到松属的共祖先图谱上。分析中共发现了6个LOD支持度强的SDL位点。这些位点分布于4个不同的连锁群上。由于这些位点控制作图亲本产生的配子向后代的不均等传递，SDL对应的基因组区域比基因组其它区域在黄山松与马尾松物种分化过程中应起着更为重要的作用。SDL分析是在远缘杂交组合中解析控制配子不均等传递遗传因子重要环节，这部分工作为SDL分析提供了一个研究实例，也是广大演化生物学家普遍感兴趣的一个研究内容。
Speciation is one of the most interesting processes in evolution, and reproductive isolationis essential for the process of speciation to be complete. Thus, understanding the geneticmechanism triggering reproductive isolation is a key goal in the study of speciation. To date, theunderlying causes of reproductive isolation are only partially understood in a few species.Hybridization between different species often has negative consequences that result in reducedfertility or viability, which leads to transmission ratio distortion (TRD) of the parental gametes.Occasionally, hybridization can raise hybrid vigor in which a hybrid offspring performs well.Yet, recombination and allelic segregation may render hybrid breakdown in the next generation.Genetic mapping methods provide a unique opportunity to study the interactions ofdifferentiated genes and genomes in a hybrid genetic background. The degree of TRD isthought to be positively correlated with the level of genome divergence between taxa, mappingthe SDLs that control the TRD marker clusters can be used to infer the genome regions relatingto reproductive barriers in distant crosses. To date, the majority evidences of TRD occurring inhybrid genetic backgrounds are based on studies employing experimental hybrids. By contrast,map-based studies of natural hybrids are much rarer.
Pinus hwangshanensis and P. massoniana are native conifers in south of China. In theareas to the south of Yangzi River, P hwangshanensis is mainly found in the elevation range of800-1800m, and P. massoniana distributes below elevation of800-900m. The major differencein the ecological niches of the two species is elevation. With an increase in elevation,environmental factors, such as oxygen partial pressure, air temperature and moisture regime,soil temperature and water regime, sunray and ultraviolet light intensity, will change. Theseenvironmental factors are closely related to plant growth and fitness. They are environmentalstresses to cause differentiation in plant phenology and fitness, subsequently, to maintain thespecies-specific characteristics of the alternate species. The distribution ranges of P.hwangshanensis and P. massoniana are frequently found to be adjacent to each other, andoverlapped with a narrow contact zone. This distribution pattern leads them to be the desirableresearch materials to uncover the genetic mechanism triggering the parapatric speciationprocess. In my thesis, studies were carried out in the following aspects: Variation in conemorphology and seed characteristics for P.massoniana, P. Hwangshanensis, and their hybridzone; Variation trends of physiological and biochemical indexes of P. hwangshanensis withincrease in elevation; AFLP protocol optimization and primer combination screening for Pinusspp.; Potential chromosomal introgression barriers revealed by linkage analysis in a hybrid of P. massoniana and P. hwangshanensis; Genome-wide detection of genetic loci triggering unevendescending of gametes from a natural hybrid pine. The main context include:
1. Variation in cone morphology and seed characteristics for P. massoniana, P.hwangshanensis and their hybrid zone. The cones and seeds were collected from differentelevations of Mt. Huang in Anhui Province. Six characteristics of cones and seeds, includingcone length, cone width, ratio of cone length to width, seed plumpness, germinationratio and weight per thousand seeds were analyzed to reveal the variation trends with thechanging of elevations. The results indicated that: Cone length, cone width, ratio of conelength to width decreased with increase in elevation; Seed plumpness, germination ratioand weight per thousand seeds were lower for plants in hybrid zone than that of theother zones. We propose it may relate to the reproduction barrier between P. massonianaand P. hwangshanensis. It was noticeable that the majority of variation for cones andseeds arose among individual trees, and traits were commonly diverged amongindividuals at different elevations.
2. Variation trends of physiological and biochemical indexes of P. hwangshanensiswith increase in elevation. P. hwangshanensis seeds collected from different elevations ofHuanggang Mountain in Wuyi Nature Reserve were cultivated in a container at25℃. Aftersprouting，the seedlings were stored in a refrigerator within a temperature range of2-5℃for24hours. Five physiological and biochemical indices of P. hwangshanensis seedlings weredetermined before and after the treatment，and the variation trends of these indexes withincreasing elevation were also studied. Three of the tested indices，including relative electricalconductivity of extract fluid，SOD activity，and MDA content of the seedlings after lowtemperature treatment decreased with increasing elevation. Therefore，the above three indicesare related to the adaptability of P. hwangshanensis to elevation. These indices could be appliedto explore the mechanism of parapatric speciation.
3. AFLP protocol optimization and primer combinations screening for Pinus spp.Pines possess gigantic genomes (10-20Gb), which are about10fold that of the human genomeand about40fold that of the poplar genome, leading them out of the genome range suitable forthe common AFLP(Amplified Fragment Length Polymorphism) protocol. This paper speciallyoptimized an economic and reliable AFLP analysis protocol for pines. The number of selectivenucleotides added to the core primers and the primer combinations suitable for AFLP analysisin such species were tested. Some AFLP primer combinations that were screened their utilityacross different species were tested. The results could also be used as a reference for allowingbetter amplification in complex templates isolated from plants species with large genomes
4. Potential chromosomal introgression barriers revealed by linkage analysis in ahybrid of P. massoniana and P. hwangshanensis. Distortion of marker transmission ratio isfrequently ascribed to selection against alleles that cause hybrid incompatibility. The naturalintrogression between P. massoniana and P. hwangshanensis and their distribution ranges lead to the emergence of the two species as desirable organisms to study the genetic mechanisms forspeciation. Using seeds sampled from trees at different elevations, we consistently detectedsharp decreases in seed germination rates of trees in the hybrid zone, which might be duelargely to the hybrid incompatibility. A genetic map was established using192megagametophytes from a single tree in the hybrid zone of the two species. Segregationdistortion analysis revealed that the percentage of significant-segregation-distortion (SSD)markers was extremely high, accounting for more than25%of the segregating markers. Theextension range, the distortion direction, and the distortion intensity of SSD markers also varieddramatically on different linkage groups. This study displays the potential chromosomalintrogression barriers between P. massoniana and P. hwangshanensis. The established mapprovides a valuable platform for conducting genome-wide association of hybrid incompatibleQTLs and/or candidate genes with marker transmission ratio distortion in the hybrid.
5. Genome-wide detection of genetic loci triggering uneven descending of gametesfrom a natural hybrid pine. Marker transmission ratio distortion (TRD) revealed in geneticmapping studies of distant crosses can be used to infer the genetic basis relating to reproductivebarriers between species. Unlike measure the degree of TRD by the overall number ofsegregation distorted markers in the affected genome regions, mapping the segregationdistorting loci (SDL) provides reliable statistic parameters that help to confine the targetgenomic region for further characterization at molecular level. Using the linkage mapconstructed for a natural hybrid of P. hwangshanensis and P. massoniana, we performed SDLanalyses and aligned the established map to the pine consensus map. Altogether,6SDLs withrelatively strong LOD supports were detected on4linkage groups of the established map. Sincegametes inheriting different alternate chromatid blocks from the SDL affecting genome regionshad uneven chance to descend to the offsprings, the corresponding genome regions weresupposed to play more significant role in rendering the speciation of P. hwangshanensis and P.massoniana. This paper presents a case study on a crucial step for uncovering the geneticfactors that triggered the uneven descending of gametes in a natural distant cross, and it maydraw broad interest for the evolutionary biologists.
引文
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