山茶属有木栓瘤植物叶片形态解剖、FTIR及其分类学意义
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摘要
山茶属(Camellia)是山茶科(Theaceae)最大的一个属,也是在系统进化上较原始的一个属。无论对于亚属(subgenus)、组(section)还是种(species)的划分,山茶属植物在分类学上都争议较大。Sealy分类系统、张宏达分类系统以及闵天禄分类系统为世界著名的三大山茶属分类系统,但是这三大分类系统之间存在巨大的分歧。因此,进一步研究山茶的分类仍然具有重要意义。
本课题研究了山茶属(genus Camellia L.)有木栓瘤(cork wart)植物金花茶组(sect.Chrysanthae Chang)、古茶组(sect.ArchecamelliaSealy)、实果茶组[sect.Stereocarpus(Pierre)Sealy]以及糙果茶组(sect.Furfuracea Chang)的叶形态解剖特征、FTIR(Fourier TransformInfrared Spectroscopy)特征以及叶片木栓瘤结构的发育解剖,为这四个组的分类提供了新的数据,澄清了许多山茶分类问题,对山茶属植物的系统分类有了新的认识。
我们的研究结果表明:
(1)山茶属植物叶形丰富多样,难以用传统的方法准确描述。叶形扫描以后,得到的长度、宽度、宽长比、周长、叶面积以及叶形系数标准化处理以后,作主成分分析。结果表明,糙果茶组植物叶形大而阔;实果茶组植物叶面积最小;古茶组和金花茶组植物叶形相似,但是经过叶形数据主成分分析,发现金花茶组植物和古茶组植物叶形仍然存在明显的差异。总的说来,金花茶组、古茶组、实果茶组以及糙果茶组的叶形存在明显差异,可以比较明显地区分开来,从而支持张宏达将这四个组作为独立的组。
(2)山茶属植物的表皮特征复杂多样,不同类群之间的叶表皮形态存在明显差异,但是在同一类群叶表皮往往有许多相似而且稳定的特征,因而对于属下分类群——组与种的划分具有重要意义。古茶组上表皮细胞异形(两种形态的表皮细胞交错分布);金花茶组上表皮细胞有晶体;糙果茶组上表皮细胞形态一般为多边形。形态解剖数据聚类分析的结果严格支持张宏达关于组的划分。
(3)各种山茶属植物由于叶片化学成分的差异因而在FTIR谱图上表现波峰和波数的差异。四组山茶属植物叶片FTIR光谱差异主要存在于碳水化合物指纹区(carbohydrate fingerprinting region)(波数wavenumbers范围1500 cm~(-1)-1000 cm~(-1))。FTIR数据聚类分析将四个组的植物在组的水平上聚成三枝,糙果茶组和金花茶组植物分别组成一枝,古茶组和实果茶组植物组成一枝,反映古茶组、金花茶组、糙果茶组以及实果茶组亲缘关系。FTIR数据支持张宏达将金花茶组、糙果茶组作为独立的组,微弱支持古茶组和实果茶组合并,但仍然有待于更深一步进行研究。
(4)木栓瘤是由早期叶片上的大型气孔(D型气孔)发育来的。D型气孔发生在山茶属植物叶片发育的早期(叶片还未展开),分布频率低。典型气孔出现在叶片展开以后,分布频率高。叶片成熟过程中,D型气孔的副卫细胞继续分裂并逐渐木栓化,形成木栓瘤。压力实验表明,木栓瘤具有气体交换的作用。木栓瘤非常有可能是后生结构,这种结构对于增加叶片的气体交换具有重要意义。但是,增加气体交换的同时可能导致水分大量散失。所以这些植物不适于生活在干旱地区,而更适合于生活于雨量充沛的地区。更深入地研究山茶属有木栓瘤植物幼叶中两类气孔的生理功能将有助于揭示山茶属植物进化与分布。
综上所述,山茶属叶形态解剖特征、FTIR数据支持张宏达山茶分类系统关于金花茶组、古茶组、实果茶组以及糙果茶组的划分。同时,本研究也为进一步研究山茶属植物分类打下了基础。
Camellia L.is the largest but relatively original genus in family Theaceae.The taxonomic treatments of the genus Camellia are still bitterly controversial in classifications of subgenera,sections or species. There are three recent taxonomic treatments of the genus Camellia frequently referred to in Camellia literatures:Sealy's treatment,Chang's treatment,Ming's treatment,but these treatments are still controversial. Therefore,there still are significant to further study the taxonomic problem of genus Camellia.
The leaf FTIR,morphology,anatomy,and the development of cork wart of Camellia plants,including sect.Archecamellia Sealy,sect. Stereocarpus(Pierre)Sealy,sect.Furfuracea Chang and sect. Chrysanthae Chang,were investigated in this study.These investigations had clarified many Camellia taxonomic problems and contributed to understand the phylogenesis of genus Camellia.
Based on our researches,we concluded as followings:
(ⅰ)A simple definition of leaf shape had been proved to be a difficult exercise,due to the extreme diversity.The PCA were conducted using the transformed value of the measurements of area,perimeter, width,length,aspect ratio(width/length)and leaf formcoefficient obtained by scanning leaf shape.Large and broad leaf shapes were specific to sect.Chrysanthae.The leaves of sect.Stereocarpus were small.We found that the leaf shapes were different among sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea,sect. Chrysanthae.These samples were obviously clustered into four groups and therefore supported Chang's treatment.
(ⅱ)The epidermal characters were various among different taxa and constant within the same taxon,and therefore they were significant to the classifications of taxa below genus:section and species. Various epidermal cells were found in sect.Archecamellia(there were two different epidermal cells arranging alternately).There were crystals in epidermal cells of sect.Chrysanthae.Most of epidermal cells in sect.Furfuracea were polygonal.
(ⅲ)Different leaf chemical compositions among Camellia species resulted in various peaks and wavenumbers in the FTIR spectrum. The comparisons of leaf FTIR indicated that most differences were specific to the carbohydrate fingerprinting region(wavenumbers 1500cm~(-1)-1000cm~(-1)).FTIR characters figured out the relationships among sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea, sect.Chrysanthae.FTIR data supported Chang treated sect. Furfuracea,sect.Chrysanthae as natural taxa.It seems that sect. Archecamellia and sect.Stereocarpus should been merged,but this need further research.
(ⅳ)Cork wart originated from D-type stoma which was large, infrequent and emerged on young leaf at early developmental stages.Typical stoma was small,frequent and emerged on young, unfolded leaf.The subsidial7 cells of D-type stoma divided and suberized gradually in the progress of maturing.Cork wart finally formed.Pressure experiment indicated that cork wart played a role of gas exchange.Cork wart is probably evolutionary character which contributes to gas exchange.However,it also increases water loss.Therefore,these plants usually distribute in areas of abundant rainfall while arid or semiarid environment is not suitable to them.Further researches about the physiological functions of the two types of stomata in young leaves of Camellia plants with cork wart will contribute to understand the evolution and distribution of genus Camellia.
To sum up,Chang's taxonomic treatment about the classifications of sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea and sect. Chrysanthae was supported by the data of leaf FTIR,morphology and anatomy.Additionally,this study laid a base for further researches on Camellia taxonomy.
本课题研究了山茶属(genus Camellia L.)有木栓瘤(cork wart)植物金花茶组(sect.Chrysanthae Chang)、古茶组(sect.ArchecamelliaSealy)、实果茶组[sect.Stereocarpus(Pierre)Sealy]以及糙果茶组(sect.Furfuracea Chang)的叶形态解剖特征、FTIR(Fourier TransformInfrared Spectroscopy)特征以及叶片木栓瘤结构的发育解剖,为这四个组的分类提供了新的数据,澄清了许多山茶分类问题,对山茶属植物的系统分类有了新的认识。
我们的研究结果表明:
(1)山茶属植物叶形丰富多样,难以用传统的方法准确描述。叶形扫描以后,得到的长度、宽度、宽长比、周长、叶面积以及叶形系数标准化处理以后,作主成分分析。结果表明,糙果茶组植物叶形大而阔;实果茶组植物叶面积最小;古茶组和金花茶组植物叶形相似,但是经过叶形数据主成分分析,发现金花茶组植物和古茶组植物叶形仍然存在明显的差异。总的说来,金花茶组、古茶组、实果茶组以及糙果茶组的叶形存在明显差异,可以比较明显地区分开来,从而支持张宏达将这四个组作为独立的组。
(2)山茶属植物的表皮特征复杂多样,不同类群之间的叶表皮形态存在明显差异,但是在同一类群叶表皮往往有许多相似而且稳定的特征,因而对于属下分类群——组与种的划分具有重要意义。古茶组上表皮细胞异形(两种形态的表皮细胞交错分布);金花茶组上表皮细胞有晶体;糙果茶组上表皮细胞形态一般为多边形。形态解剖数据聚类分析的结果严格支持张宏达关于组的划分。
(3)各种山茶属植物由于叶片化学成分的差异因而在FTIR谱图上表现波峰和波数的差异。四组山茶属植物叶片FTIR光谱差异主要存在于碳水化合物指纹区(carbohydrate fingerprinting region)(波数wavenumbers范围1500 cm~(-1)-1000 cm~(-1))。FTIR数据聚类分析将四个组的植物在组的水平上聚成三枝,糙果茶组和金花茶组植物分别组成一枝,古茶组和实果茶组植物组成一枝,反映古茶组、金花茶组、糙果茶组以及实果茶组亲缘关系。FTIR数据支持张宏达将金花茶组、糙果茶组作为独立的组,微弱支持古茶组和实果茶组合并,但仍然有待于更深一步进行研究。
(4)木栓瘤是由早期叶片上的大型气孔(D型气孔)发育来的。D型气孔发生在山茶属植物叶片发育的早期(叶片还未展开),分布频率低。典型气孔出现在叶片展开以后,分布频率高。叶片成熟过程中,D型气孔的副卫细胞继续分裂并逐渐木栓化,形成木栓瘤。压力实验表明,木栓瘤具有气体交换的作用。木栓瘤非常有可能是后生结构,这种结构对于增加叶片的气体交换具有重要意义。但是,增加气体交换的同时可能导致水分大量散失。所以这些植物不适于生活在干旱地区,而更适合于生活于雨量充沛的地区。更深入地研究山茶属有木栓瘤植物幼叶中两类气孔的生理功能将有助于揭示山茶属植物进化与分布。
综上所述,山茶属叶形态解剖特征、FTIR数据支持张宏达山茶分类系统关于金花茶组、古茶组、实果茶组以及糙果茶组的划分。同时,本研究也为进一步研究山茶属植物分类打下了基础。
Camellia L.is the largest but relatively original genus in family Theaceae.The taxonomic treatments of the genus Camellia are still bitterly controversial in classifications of subgenera,sections or species. There are three recent taxonomic treatments of the genus Camellia frequently referred to in Camellia literatures:Sealy's treatment,Chang's treatment,Ming's treatment,but these treatments are still controversial. Therefore,there still are significant to further study the taxonomic problem of genus Camellia.
The leaf FTIR,morphology,anatomy,and the development of cork wart of Camellia plants,including sect.Archecamellia Sealy,sect. Stereocarpus(Pierre)Sealy,sect.Furfuracea Chang and sect. Chrysanthae Chang,were investigated in this study.These investigations had clarified many Camellia taxonomic problems and contributed to understand the phylogenesis of genus Camellia.
Based on our researches,we concluded as followings:
(ⅰ)A simple definition of leaf shape had been proved to be a difficult exercise,due to the extreme diversity.The PCA were conducted using the transformed value of the measurements of area,perimeter, width,length,aspect ratio(width/length)and leaf formcoefficient obtained by scanning leaf shape.Large and broad leaf shapes were specific to sect.Chrysanthae.The leaves of sect.Stereocarpus were small.We found that the leaf shapes were different among sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea,sect. Chrysanthae.These samples were obviously clustered into four groups and therefore supported Chang's treatment.
(ⅱ)The epidermal characters were various among different taxa and constant within the same taxon,and therefore they were significant to the classifications of taxa below genus:section and species. Various epidermal cells were found in sect.Archecamellia(there were two different epidermal cells arranging alternately).There were crystals in epidermal cells of sect.Chrysanthae.Most of epidermal cells in sect.Furfuracea were polygonal.
(ⅲ)Different leaf chemical compositions among Camellia species resulted in various peaks and wavenumbers in the FTIR spectrum. The comparisons of leaf FTIR indicated that most differences were specific to the carbohydrate fingerprinting region(wavenumbers 1500cm~(-1)-1000cm~(-1)).FTIR characters figured out the relationships among sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea, sect.Chrysanthae.FTIR data supported Chang treated sect. Furfuracea,sect.Chrysanthae as natural taxa.It seems that sect. Archecamellia and sect.Stereocarpus should been merged,but this need further research.
(ⅳ)Cork wart originated from D-type stoma which was large, infrequent and emerged on young leaf at early developmental stages.Typical stoma was small,frequent and emerged on young, unfolded leaf.The subsidial7 cells of D-type stoma divided and suberized gradually in the progress of maturing.Cork wart finally formed.Pressure experiment indicated that cork wart played a role of gas exchange.Cork wart is probably evolutionary character which contributes to gas exchange.However,it also increases water loss.Therefore,these plants usually distribute in areas of abundant rainfall while arid or semiarid environment is not suitable to them.Further researches about the physiological functions of the two types of stomata in young leaves of Camellia plants with cork wart will contribute to understand the evolution and distribution of genus Camellia.
To sum up,Chang's taxonomic treatment about the classifications of sect.Archecamellia,sect.Stereocarpus,sect.Furfuracea and sect. Chrysanthae was supported by the data of leaf FTIR,morphology and anatomy.Additionally,this study laid a base for further researches on Camellia taxonomy.
引文
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