高原芥三个居群物种生物学特性及十字花科部分属种的孢粉学研究
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摘要
以帕米尔高原地区的高原芥(Christolea crassifolia Camb.)为研究对象,应用居群生物学的原理和方法,对高原芥三个不同居群的核型、花粉形态、叶表皮气孔形态、表型特征进行全面的研究;并对南芥族(Arabideae)、紫罗兰族(Matthioleae)、香花芥族(Hesperideae)和大蒜芥族(Sisymbrieae)4族8属13种(包括3个居群高原芥)植物的花粉形态进行观察,结果显示:
(1)高原芥的三个居群
①卡拉苏居群核型公式为K=2n=2x=14=10m+2sm(SAT)+2sm,布伦口居群核型公式为K=2n=2x=14=6m+2m(SAT)+6sm,瓦恰居群核型公式为K=2n=2x=14=12m+2sm(SAT),核型类型均为1A型;核型不对称系数分别为卡拉苏居群60.30%,布伦口居群60.86%,瓦恰居群60.17%。
②通过扫描电镜观察发现,高原芥3个居群的花粉均具三沟萌发孔,沟长达两极,极面观都为三裂圆形,网状纹饰,但在花粉形状,网眼直径和网脊宽度上存在差异。
③高原芥三个居群叶上、下表皮细胞形状均为3～6边组成的多边型,垂周壁平直,上下表皮均有气孔分布。布伦口和瓦恰居群上表皮气孔密度小于下表皮气孔密度,而卡拉苏居群上表皮气孔密度大于下表皮,三个居群气孔类型均为无规则型,保卫细胞肾形,无副卫细胞。
④高原芥三个居群32个形态性状UPGMA聚类分析的结果显示,卡拉苏居群单独构成一支,而布伦口和瓦恰居群构成一支。对高原芥三个居群的32个性状进行了方差分析,其中25个性状差异显著,7个性状差异不显著。
⑤高原芥的染色体数目、核型不对称系数、花粉萌发孔类型、花粉极面观、花粉外壁纹饰、叶上、下表皮细胞形状、垂周壁式样、叶上气孔分布位置、气孔类型、保卫细胞形状、叶形系数、花瓣长、花瓣宽、雄蕊花丝长度,这些性状特征稳定性较强,不同居群间没有明显差异。
(2)花粉形态
4族8属13种植物花粉的一般特征为:花粉粒长球形至超长球形,外壁纹饰为网状或细网状,三沟或三、四沟,并根据花粉形态、外壁纹饰、网眼大小和萌发沟数目等特征将所研究的13种十字花科植物的花粉形态特征分为5种类型:①花粉粒长球形,网状纹饰,三沟;②花粉粒超长球形,网状纹饰,三沟;③花粉粒长球形,细网状纹饰,三沟;④花粉粒超长球形,细网状纹饰,三沟;⑤花粉粒长球形,三或四沟,网状纹饰。
The karyotype,pollen morphology,stoma traits of epidermis,and exterior appearance of Christolea crassifolia in three different populations of the Parmirs have been investigated completely by the principles and methods of populations in this paper.Pollen morphology of 13 species(including Ch.crassifolia in three populations) belonging to 8 genera of 4 tribes(Arabideae,Matthioleae,Hesperideae,Sisymbrieae) were studied.The results indicated that:
(1) Three populations of Ch.crassifolia
①The karyotype formula of Kalasu population is:K=2n=2x=14=10m+2sm(SAT)+2sm,the karyotype formula of Bulunkou population is:K=2n=2x=14=6m+2m(SAT)+6sm,the karyotype formula of Waqia population is:K=2n=2x=14=12m+2sm(SAT).The karyotypes of the 3 populations are all Stebbins's 1A type,and the asymmetric coefficients are Kalasu's 60.30%,Bulunkou's 60.86%,Waqia's 60.17%.
②The observations of scanning electron microscope showed that the pollens of the three poulations are all three-colpate,with reticulate ornamentation and the polar views are all three-lobed circle,but there are significantly differences in the shape of pollen,the diameter of areole and the width of murus.
③The shape of upper and lower epidermis cells of Ch.crassifolia in three populations are all polygons with 3～6 sides,the anticlinal walls are straight,and stomatic distributed in upper and lower epidermis.The stomatal density of upper epidermis is denser than that of lower epidermis in Kalasu population comparing with Bulunkou and Waqia populations.The stomatal apparatus of three populations are all irregular types,and the guard cells are all kidney-shaped and there are no subsidiary cell.
④The UPGMA dendrogram using 32 traits for 3 populations of Ch.Crassifolia shows that,Kalasu population formed one group alone,Bulunkou and Waqia populations formed another group.Variance analysis of 32 traits among 3 populations of Ch.crassifolia indicates that 25 traits have evident difference, and seven traits have no evident difference.
⑤The chromosome numbers,asymmetric coefficient,the type of germinal aperture,polar view,exine sculpture,the shape of upper and lower epidermis cells,anticlinal wall,the distribution of stomatal,the type of stomatal apparatus,the shape of guard cells,leaf shape index,petal length,petal width,filament length of Ch.Crassifolia have good stability.
(2) The pollen morphology
The pollen morphology of 13 species belonging to 8 genera of 4 tribes is very similar,their shape varies from prolate to perprolate with reticulate ornamentation,the pollen grains are most tricolpate(rarely teracolpate).Five pollen types can be distinguished based on the shape of pollen grains,the size of the areole and the number of the aperture:①Pollen grains three-colpate,prolate,with reticulate ornamentation;②Pollen grains three-colpate,perprolate,with reticulate ornamentation;③Pollen grains three-colpate, prolate,with microreticulate ornamentation;④Pollen grains three-colpate,perprolate,with microreticulate ornamentation;⑤Pollen grains three-colpate or four-colpate,prolate,with reticulate ornamentation.
(1)高原芥的三个居群
①卡拉苏居群核型公式为K=2n=2x=14=10m+2sm(SAT)+2sm,布伦口居群核型公式为K=2n=2x=14=6m+2m(SAT)+6sm,瓦恰居群核型公式为K=2n=2x=14=12m+2sm(SAT),核型类型均为1A型;核型不对称系数分别为卡拉苏居群60.30%,布伦口居群60.86%,瓦恰居群60.17%。
②通过扫描电镜观察发现,高原芥3个居群的花粉均具三沟萌发孔,沟长达两极,极面观都为三裂圆形,网状纹饰,但在花粉形状,网眼直径和网脊宽度上存在差异。
③高原芥三个居群叶上、下表皮细胞形状均为3～6边组成的多边型,垂周壁平直,上下表皮均有气孔分布。布伦口和瓦恰居群上表皮气孔密度小于下表皮气孔密度,而卡拉苏居群上表皮气孔密度大于下表皮,三个居群气孔类型均为无规则型,保卫细胞肾形,无副卫细胞。
④高原芥三个居群32个形态性状UPGMA聚类分析的结果显示,卡拉苏居群单独构成一支,而布伦口和瓦恰居群构成一支。对高原芥三个居群的32个性状进行了方差分析,其中25个性状差异显著,7个性状差异不显著。
⑤高原芥的染色体数目、核型不对称系数、花粉萌发孔类型、花粉极面观、花粉外壁纹饰、叶上、下表皮细胞形状、垂周壁式样、叶上气孔分布位置、气孔类型、保卫细胞形状、叶形系数、花瓣长、花瓣宽、雄蕊花丝长度,这些性状特征稳定性较强,不同居群间没有明显差异。
(2)花粉形态
4族8属13种植物花粉的一般特征为:花粉粒长球形至超长球形,外壁纹饰为网状或细网状,三沟或三、四沟,并根据花粉形态、外壁纹饰、网眼大小和萌发沟数目等特征将所研究的13种十字花科植物的花粉形态特征分为5种类型:①花粉粒长球形,网状纹饰,三沟;②花粉粒超长球形,网状纹饰,三沟;③花粉粒长球形,细网状纹饰,三沟;④花粉粒超长球形,细网状纹饰,三沟;⑤花粉粒长球形,三或四沟,网状纹饰。
The karyotype,pollen morphology,stoma traits of epidermis,and exterior appearance of Christolea crassifolia in three different populations of the Parmirs have been investigated completely by the principles and methods of populations in this paper.Pollen morphology of 13 species(including Ch.crassifolia in three populations) belonging to 8 genera of 4 tribes(Arabideae,Matthioleae,Hesperideae,Sisymbrieae) were studied.The results indicated that:
(1) Three populations of Ch.crassifolia
①The karyotype formula of Kalasu population is:K=2n=2x=14=10m+2sm(SAT)+2sm,the karyotype formula of Bulunkou population is:K=2n=2x=14=6m+2m(SAT)+6sm,the karyotype formula of Waqia population is:K=2n=2x=14=12m+2sm(SAT).The karyotypes of the 3 populations are all Stebbins's 1A type,and the asymmetric coefficients are Kalasu's 60.30%,Bulunkou's 60.86%,Waqia's 60.17%.
②The observations of scanning electron microscope showed that the pollens of the three poulations are all three-colpate,with reticulate ornamentation and the polar views are all three-lobed circle,but there are significantly differences in the shape of pollen,the diameter of areole and the width of murus.
③The shape of upper and lower epidermis cells of Ch.crassifolia in three populations are all polygons with 3～6 sides,the anticlinal walls are straight,and stomatic distributed in upper and lower epidermis.The stomatal density of upper epidermis is denser than that of lower epidermis in Kalasu population comparing with Bulunkou and Waqia populations.The stomatal apparatus of three populations are all irregular types,and the guard cells are all kidney-shaped and there are no subsidiary cell.
④The UPGMA dendrogram using 32 traits for 3 populations of Ch.Crassifolia shows that,Kalasu population formed one group alone,Bulunkou and Waqia populations formed another group.Variance analysis of 32 traits among 3 populations of Ch.crassifolia indicates that 25 traits have evident difference, and seven traits have no evident difference.
⑤The chromosome numbers,asymmetric coefficient,the type of germinal aperture,polar view,exine sculpture,the shape of upper and lower epidermis cells,anticlinal wall,the distribution of stomatal,the type of stomatal apparatus,the shape of guard cells,leaf shape index,petal length,petal width,filament length of Ch.Crassifolia have good stability.
(2) The pollen morphology
The pollen morphology of 13 species belonging to 8 genera of 4 tribes is very similar,their shape varies from prolate to perprolate with reticulate ornamentation,the pollen grains are most tricolpate(rarely teracolpate).Five pollen types can be distinguished based on the shape of pollen grains,the size of the areole and the number of the aperture:①Pollen grains three-colpate,prolate,with reticulate ornamentation;②Pollen grains three-colpate,perprolate,with reticulate ornamentation;③Pollen grains three-colpate, prolate,with microreticulate ornamentation;④Pollen grains three-colpate,perprolate,with microreticulate ornamentation;⑤Pollen grains three-colpate or four-colpate,prolate,with reticulate ornamentation.
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