论文摘要
本文研究了广义蓼属(Polygonum L.)以及相关属(Fallopia,Reynoutria,Antenoron,Fagopyrum和Pteroxygonum)的分子系统学、果实形态解剖学、花被片形态学。在此基础上,对中国的单型属Pteroxygonum进行了系统学位置的研究,主要研究结果如下:果实形态解剖学研究结果表明,所观察的物种中外果皮常常加厚,并且可以根据外果皮细胞的垂周壁是否平直,可以明显的区分为平直型和波状两类,具有平直型垂周壁的外果皮细胞的主要特征是细胞为方形或者为高度为宽度的1.5-2倍的矩形,并拥有二叉状分枝的内腔,此种类型的我们所观察到的植物有Polygonum sect.Aconogonon和Polygonum sect.Bistorta。具有波状型垂周壁的外果皮细胞的主要特征是细胞为矩形,其高度为宽度的2倍-4倍,内腔为直的不分叉或者没有内腔,此种类型的我们所观察到的植物有sect.Persicaria,Antenoron,Polygonum sect.Echinocaulon和Polygonum sect.Cephalophilon。Fagopyrum具有较为特殊的外果皮形态,一种为外果皮不加厚,另外一种为外果皮加厚并有内腔分布。其他的类群没有发现一致的解剖学性状。基于核糖体基因5.8S以及间隔片断ITS的序列分析,对36种蓼属及相关属进行了系统演化关系分析。研究结果表明由Fallopia,Reynoutria和Polygonum sect.Polygonum构成的Polygoneae(Ⅰ)这一支,与Fagopyrum,Koenigia,Polygonum sect.Bistorta,Polygonum sect.Cephalophilon,Polygonum sect.Persicaria,Antenoron以及Pteroxygonum组成的Persicarieae(Ⅱ)一支构成姐妹群。在Persicarieae中,Fagopyrum是最早分化的一个谱系,Pteroxygonum紧随其后分化出来,二者依次和Polygonum(except Polygonum sect.Polygonum)构成姐妹类群关系。在Polygonum(except Polygonum sect.Polygonum)这一支中分化出两大亚分支:一支是由Polygonum sect.Bistorta,Polygonum sect.Aconogonon和Koenigia组成,其中Polygonum sect.Aconogonon和Koenigia聚合成的B这一支与Polygonum sect.Bistorta的A这一支形成姐妹关系;两外一支是由Polygonum sect.Cephalophilon,Polygonum sect.Persicaria,Polygonum sect.Echinocaulon和Antenoron组成的,但是他们之间的关系并没有得到很高的支持率的支持。基于ITS的系统发育树的研究支持将蓼属于及相关属划分为个族来处理即tribes Persicarieae和Polygoneae。结合前人的观点,我们支持Koenigia是Polygonum sect.Aconogonon在高原地区的进化类型,并且承认Aconogonon属级位置的存在。针对Koenigia的物种范围,ITS分子系统同样选取了蓝药蓼Polygonum cyanandrum和大铜钱叶蓼Polygonum forrestii作为探讨Koenigia范围的物种,结果同atpB-rbcL所得到的结果一致即这两个种应置于冰岛蓼属,支持将这个类群作为一个属处理。根据ITS系统发育图,Polygonum sect.Bistorta这个类群植物与Polygonum sect.Aconogonon的亲缘关系较近即(A+B)这一支,建议将这两个类群的分类阶元上升为属,同时ITS系统发育树的结果表明Fagopyrum置于tribe Persicarieae中,并且位于基部位置,同意Ronse Decreane的观点即Fagopyrum是tribe Persicarieae的基部类群,不赞同Fagopyrum与Fallopia具有较近亲缘关系的观点。针对广义蓼属的范围问题,我们根据分子系统学的证据,认为应该成立Persicaria其下包括以下的几个组:sects.Echinocaulon,Cephalophilon,Persicaria,Tovara。本文特别讨论了Polygonum bungeanum的归属问题,根据形态学特征以及分子系统学的结果认为Polygonum bungeanum应该属于sect.Persicaria。在我们的ITS系统发育树上,由于I group中共包括了Fallopia和sect.Polygonum两个类群,也许是由于我们的分子生物学试验中sect.Polygonum的物种选择不多,所以导致了Polygonum sect.Polygonum插入到Fallopia支中。I group中Fallopia与Reynoutria聚为一支,所以我们更倾向于将Reynoutria作为Fallopia的一个组来处理,同时我们发现在I group中除了有些基部有支持率比较高的区分外,内部的各分支的自展支持率并不高,从而反映出这个属的分组状态从我们的ITS系统发育图中并不能很好的反映出来,有必要通过更多的物种和基因片断联合分析这个属的分组情况。通过对翼蓼及相关属(蓼属、首乌属、虎杖属、荞麦属和金线草属)的形态观察、果实解剖学观察、花被片脉序观察、花粉形态、核型分析、以及ITS序列的分析确定了翼蓼和荞麦较远的亲缘关系。其中我们发现翼蓼果实基部有三个角物明显不同于其他属果实的形态特征。翼蓼外果皮明显加厚并有零星散布的波状内腔,而荞麦的外果皮很薄细胞不等径中果皮极厚。以上证据证明了翼蓼与荞麦属有较远亲缘关系。在观察荞麦属和翼蓼的花被片脉络时发现了两种不同的脉序类型,符合将荞麦属被分为两个组的划分。翼蓼花被片脉序为三出状,支持将翼蓼归为Persicarieae族。对翼蓼及荞麦属植的花粉进行了比较后,发现荞麦属植物的花粉网孔有明显的内凹穿孔而翼蓼却没有,结果表明二者较远的亲缘关系。通过对nrDNA ITS区域序列分析得出翼蓼及相关属为一个单系类群,含有两个稳定的分支:第一个分支由蓼属(萹蓄组)、首乌属、虎杖属的植物组成,第二个分支由蓼属(刺蓼组、蓼组、分叉蓼组、拳参组)、翼蓼和荞麦属植物组成。同时第二个分支又分成了两个亚分支,蓼属(刺蓼组、蓼组、分叉蓼组、拳参组)和翼蓼属植物属于第一个亚支而荞麦属植物植物属于第二个亚支。结果支持翼蓼不属于荞麦属的范畴。实验结果显示翼蓼是个单型属,并且属于Persicarieae族。
论文目录
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