马褂木遗传多样性及系统地理学研究
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摘要
马褂木(鹅掌楸)(Liriodendron chinense (Hemsl.) Sarg.)是我国特有的珍稀濒危第三纪孑遗树种,也是典型的东亚-北美间断分布种,对古植物学、植物系统学和植物地理学研究具有重要的科学价值；马褂木也是优良的用材、绿化和园林观赏树种,具有重要的经济价值和生态价值。本研究以马褂木全分布区有代表性的群体为材料,从叶、花器官、花粉、种实的形态、光合特性、种子与花粉活力、核DNA及叶绿体DNA等方面进行了遗传多样性的研究,并对其系统地理学进行了探讨。主要结果如下：
1.表型水平上,马褂木的种实性状、叶型、光合特性、花器官均存在显著的群体变异,种实性状存在显著的海拔梯度变异规律,聚合果宽度随海拔的升高有变窄的趋势；叶型及花器官大小呈现出显著的径向地理变异规律,东部群体叶片侧裂凹陷及前裂凹陷程度均比西部群体明显,东部群体花朵明显小于西部；净光合速率最高的群体为最低的2.36倍；群体间显著分化。
2.花粉在群体间存在明显的多态性。花粉形态、大小、外壁纹饰及超微结构均存在显著的经向地理变异,西部花粉类型相对较为原始,东部则较为进化,东西部之间存在渐变进化类型,马褂木花粉在木兰科中属于较为进化的类型。
3.马褂木的遗传多样性处于较低水平(Nei(h)=0.5577),一定程度上反映了其濒危现状；西部和中南部群体的遗传多样性有高于东部群体的趋势,东、西部群体间存在高水平地理分化(Fst=0.3018)。其群体分化较大的原因可能为：星散岛状分布的地理隔离效应,加上大部分群体规模很小,限制了群体间的基因流,产生了明显的遗传漂变。各群体均存在显著的连锁不平衡,研究表明不平衡主要是一些随机漂变或人为干扰等因素引起的。
4.马褂木cpDNA单倍型呈显著的东、西部地理分布格局。东部以福建与浙江交界的武夷山北麓的中山地带为单倍型分布中心,西部以重庆、贵州、湖南、湖北交界的中山地带为单倍型分布中心。多数单倍型为东、西部的区域内共有,推测东、西部的马褂木可能起源于不同的避难所。西部区域的避难所可能为云贵高原和四川南部,东部区域的避难所可能为武夷山脉南麓。冰期后,推测东、西部避难所马褂木可能向北方的不同方向扩散,在重庆、贵州、湖南、湖北交界的地带交汇,导致该地区存在有丰富的单倍型。
5.无论在形态还是分子水平上,马褂木遗传变异均存在显著的东、西部地理分化,西部群体的遗传多样性高于东部,因此应优先对遗传多样高的地区进行原地保护,同时也可考虑迁地保护；进行杂交育种的亲本选配时,重视种群选择是提高育种效率的重要途径。
Liriodendron Chinense is an endangered deciduous perennial xylophyta, one of the genera with a East Asia-North American disjunctio. It has important scientific value for study on paleobotany, genealogy and geobotany. As a, relic species from the tertiary, mainly occurs in subtropical China. It has high economic and ecological value. In this research, the genetic diversity and phylogeography of L.Chinense were studied.
The main results as follows:
1. Morphological diversities among populations were discussed on the basis of traits such as the leaf, the seed, the aggregate fruit, the floral organs, the pollen. The above traits were significantly difference among populations. The mean germinations of most seed sources studied were at 3% to 5%. Leaf shape and floral organ size had significantly geographic variation along the longitude. Side sinus depth and top sinus depth in eastern region were deeper than in western region. Eastern flower was smaller than western's. The net photosynthetic rate of the highest provenance was 2.36 times the lowest.
2. Pollen had significant polymorphism among populations. Pollen morphology and ultrastructure were significant geographic variations. Pollen type of western region was more primitive relatively than eastern region. L. Chinense pollen was a more evolutionary type in the Magnoliaceae.
3. In this study,20 SSR primers were used to reveal the genetic diversity of 22 populations, low genetic diversity (Nei(h)=0.5577)was found. Genetic diversity has higher geographic differentiation between eastern and western (Fst=0.3018). It maybe caused by significant genetic drift which was due to geographical isolation, little gene flow and small population scale. Each population exited significant linkage disequilibria, implied that the stochastic causes such as genetic drift might be the main reason of linkage disequilibria.
4. Significant geographic structure of haplotype was detected. Populations of western and eastern ranges had own unique haplotypes, which implied that eastern and western range had different origins. Two contrasting glacial refugia were found in eastern and western distribution regions. Yunnan-Guizhou plateau and southern Sichuan maybe provided refugium for western region; Southren Wuyi-mountain maybe served the other one for eastern region. After ice age, we speculated that L.Chinenese in eastern and western refugia maybe spreaded in different directions to the north. Populations survived in eastern and western refugia migrated and then mixed in border zone intersection of Chongqing, Guizhou, Hunan and Hubei, resulting in that the region is rich in haplotypes.
5. Morphologic and molecular levels, L.Chinense, genetic variation had significant geographic disintegration in the East and West of China. The genetic diversity of western populations was higher than the eastern's. Therefore, high genetic diversity populations should have the priority in situ conservation, but also to consider the ex-situ conservation. For hybrid breeding of parent selection, the emphasis on provenance selection is an important way to improve breeding efficiency.
1.表型水平上,马褂木的种实性状、叶型、光合特性、花器官均存在显著的群体变异,种实性状存在显著的海拔梯度变异规律,聚合果宽度随海拔的升高有变窄的趋势；叶型及花器官大小呈现出显著的径向地理变异规律,东部群体叶片侧裂凹陷及前裂凹陷程度均比西部群体明显,东部群体花朵明显小于西部；净光合速率最高的群体为最低的2.36倍；群体间显著分化。
2.花粉在群体间存在明显的多态性。花粉形态、大小、外壁纹饰及超微结构均存在显著的经向地理变异,西部花粉类型相对较为原始,东部则较为进化,东西部之间存在渐变进化类型,马褂木花粉在木兰科中属于较为进化的类型。
3.马褂木的遗传多样性处于较低水平(Nei(h)=0.5577),一定程度上反映了其濒危现状；西部和中南部群体的遗传多样性有高于东部群体的趋势,东、西部群体间存在高水平地理分化(Fst=0.3018)。其群体分化较大的原因可能为：星散岛状分布的地理隔离效应,加上大部分群体规模很小,限制了群体间的基因流,产生了明显的遗传漂变。各群体均存在显著的连锁不平衡,研究表明不平衡主要是一些随机漂变或人为干扰等因素引起的。
4.马褂木cpDNA单倍型呈显著的东、西部地理分布格局。东部以福建与浙江交界的武夷山北麓的中山地带为单倍型分布中心,西部以重庆、贵州、湖南、湖北交界的中山地带为单倍型分布中心。多数单倍型为东、西部的区域内共有,推测东、西部的马褂木可能起源于不同的避难所。西部区域的避难所可能为云贵高原和四川南部,东部区域的避难所可能为武夷山脉南麓。冰期后,推测东、西部避难所马褂木可能向北方的不同方向扩散,在重庆、贵州、湖南、湖北交界的地带交汇,导致该地区存在有丰富的单倍型。
5.无论在形态还是分子水平上,马褂木遗传变异均存在显著的东、西部地理分化,西部群体的遗传多样性高于东部,因此应优先对遗传多样高的地区进行原地保护,同时也可考虑迁地保护；进行杂交育种的亲本选配时,重视种群选择是提高育种效率的重要途径。
Liriodendron Chinense is an endangered deciduous perennial xylophyta, one of the genera with a East Asia-North American disjunctio. It has important scientific value for study on paleobotany, genealogy and geobotany. As a, relic species from the tertiary, mainly occurs in subtropical China. It has high economic and ecological value. In this research, the genetic diversity and phylogeography of L.Chinense were studied.
The main results as follows:
1. Morphological diversities among populations were discussed on the basis of traits such as the leaf, the seed, the aggregate fruit, the floral organs, the pollen. The above traits were significantly difference among populations. The mean germinations of most seed sources studied were at 3% to 5%. Leaf shape and floral organ size had significantly geographic variation along the longitude. Side sinus depth and top sinus depth in eastern region were deeper than in western region. Eastern flower was smaller than western's. The net photosynthetic rate of the highest provenance was 2.36 times the lowest.
2. Pollen had significant polymorphism among populations. Pollen morphology and ultrastructure were significant geographic variations. Pollen type of western region was more primitive relatively than eastern region. L. Chinense pollen was a more evolutionary type in the Magnoliaceae.
3. In this study,20 SSR primers were used to reveal the genetic diversity of 22 populations, low genetic diversity (Nei(h)=0.5577)was found. Genetic diversity has higher geographic differentiation between eastern and western (Fst=0.3018). It maybe caused by significant genetic drift which was due to geographical isolation, little gene flow and small population scale. Each population exited significant linkage disequilibria, implied that the stochastic causes such as genetic drift might be the main reason of linkage disequilibria.
4. Significant geographic structure of haplotype was detected. Populations of western and eastern ranges had own unique haplotypes, which implied that eastern and western range had different origins. Two contrasting glacial refugia were found in eastern and western distribution regions. Yunnan-Guizhou plateau and southern Sichuan maybe provided refugium for western region; Southren Wuyi-mountain maybe served the other one for eastern region. After ice age, we speculated that L.Chinenese in eastern and western refugia maybe spreaded in different directions to the north. Populations survived in eastern and western refugia migrated and then mixed in border zone intersection of Chongqing, Guizhou, Hunan and Hubei, resulting in that the region is rich in haplotypes.
5. Morphologic and molecular levels, L.Chinense, genetic variation had significant geographic disintegration in the East and West of China. The genetic diversity of western populations was higher than the eastern's. Therefore, high genetic diversity populations should have the priority in situ conservation, but also to consider the ex-situ conservation. For hybrid breeding of parent selection, the emphasis on provenance selection is an important way to improve breeding efficiency.
引文
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