山茶属瘤果茶组系统分类及其种质资源评估
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摘要
瘤果茶组(Tuberculata)隶属山茶科(Theaceae)、山茶属(Camellia),因其“子房与果实表面具瘤状突起”而得名,也因此而被认为是山茶属内保持原始形状的特化类群。该组仅在我国西南地区发现分布,是我国亚热带地区的特有类群。目前,对瘤果茶组的系统分类存在较大的分歧,给瘤果茶组植物种质资源的保护和开发利用带来极大的不便。
本课题通过对瘤果茶组叶的微形态、叶解剖研究,并结合傅立叶红外转换光谱(FTIR)技术,对瘤果茶组现有的存在争议的18个种进行研究,为瘤果茶组植物分类研究提供新的证据,从而澄清瘤果茶组植物分类的一些问题。同时在明确原种的基础上,进一步利用叶绿素荧光技术研究瘤果茶组植物叶片叶绿素荧光特性的季节性生长变化,对瘤果茶组种质资源从光合特性的角度进行生长适应性评估;利用FTIR方法,对瘤果茶组种质资源从植物化学的角度进行功能性的评估,最终为瘤果茶组植物资源的开发和应用提供初步的理论基础和实验证据。我们的实验结果如下:
(1)被研究的18种植物材料叶上表皮细胞形态有不规则形和多边形两种,垂周壁样式为浅波、深波和直曲型3种。下表皮细胞样式比较单一,所有种的下表皮细胞形态均为不规则形,垂周壁样式均为浅波形。所有植物的叶的气孔均分布在叶下表皮,上表皮没有气孔分布。所有气孔器类型均为环列型(anisocytic type),部分叶片具有表皮毛。所有研究的植物均为异面叶,一般由蜡质层、上下表皮、栅栏组织、海绵组织和维管组织组成。叶片栅栏组织细胞层数比较稳定。大多数种被发现有复表皮的存在。所有叶肉中均含有石细胞。除冬青叶光果茶C.ilicifolia外,其它被研究植物叶肉中均发现晶体的存在。这些结果表明叶微形态解剖特征在种的鉴定上具有一定的应用价值,可以支持闵天禄系统对部分瘤果茶植物的归并问题。同时,研究也表明叶微形态解剖特征在组间的区分上表现出更明显的分类学意义,支持张宏达系统中将冬青叶光果茶从瘤果茶组中分出的观点。此外,瘤果茶组在叶表皮微形态和叶解剖特征变化较小,也表明瘤果茶组是一个亲缘关系极为接近的、自然的组。
(2)瘤果茶组植物叶FTIR数据的PCA和聚类分析表明,冬青叶光果茶和其它17种瘤果茶有明显的差异。这一结果并不支持闵天禄将冬青叶光果茶从张宏达系统的半宿萼茶组Pseudocamellia中分离出来合并至瘤果茶组中。FTIR结果也表明,不同类群和种间的聚集程度各不相同,体现了种间的差异性和相似度,部分的支持了闵天禄系统中对瘤果茶组植物进行归并的观点。
(3)基于叶表皮微形态、叶解剖和FTIR分析的瘤果茶组种的修订结果表明:瘤果茶C.tuberculata Chien,皱果茶C.rhytidocarpa Chang et,皱叶瘤果茶C.rhytidophylla Chang Liang,安龙瘤果茶C.anlungensis Chang,厚壳红瘤果茶C.rubituberculata Chang,尖苞瘤果茶C.acutiperulata Chang et Ye.,狭叶瘤果茶C.neriiflolia Chang,倒卵叶瘤果茶C.obovatifolia Chang,荔波红瘤果茶C.rubimuricataChang et Z.R.XU,小瘤果茶C.parvimuricata Chang,三江瘤果茶C.pyxidiacea Z.R.Xu et al.等11种瘤果茶被确立。同时,研究结果也支持张宏达系统将冬青叶光果茶从瘤果茶组中移入半宿萼茶组的处理。
(4)瘤果茶组植物叶片叶绿素荧光响应曲线、季节性Yield日进程及叶绿素相对含量测定的结果表明,皱叶瘤果茶、厚壳红瘤果茶、瘤果茶、皱果茶等具有较好的光能利用能力和叶绿素生长指标,因而对于本地区的环境有较强的适应性。而狭叶瘤果茶、倒卵叶瘤果茶、小瘤果茶、尖萼瘤果茶等则表现出在本地区自然条件下生长的不适应性。因此,我们认为狭叶瘤果茶、倒卵叶瘤果茶、小瘤果茶、尖苞瘤果茶等植物类群并不适合在本地区栽种,至少不适合在野外无遮荫处理栽种。同时,在本地区栽培种植瘤果茶植物过程中,适当的遮荫处理将会有助于植物更好生长。
(5)瘤果茶组植物与茶的叶FTIR光谱显示,瘤果茶植物植物叶片红外光谱在表征碳水化合物和和多糖的3400 cm~(-1),1150-1030 cm~(-1)处的吸收强度较大,这表明瘤果茶植物叶片的维管组织比茶发达。瘤果茶植物和茶在1730 cm~(-1)附近,1650 cm~(-1),1550-1510 cm~(-1),1450和1320 cm~(-1)附近,1300-1200 cm~(-1)等光谱区域有对应的吸收峰,这些吸收峰分别表征的是茶多酚、氨基酸、茶醇、羧酸、咖啡碱等活性成分,这表明瘤果茶组植物叶也含有较为丰富的活性物质,可以考虑将其作为茶多酚、氨基酸、茶醇、咖啡碱等活性物质的来源植物。
The section Tuberculata Chang, belonging to family Theacea, genus Camellia, only distributed in China, represents a good example of a taxonomic group in the genus Camellia with controversial circumscription. This section was charactered by the tuberculate presented on the ovaries and seeds, and it was therefore considered to be orignal groups in Camellia. At present, the taxonomic treatment of the section Tuberculata is still ambiguous, which causes great inconvenience to protection of the germplasm resources and the development and utilization.
In present study, the leaf epidermial micro-morphology and comparative anatomy of 18 disputed species of the section Tuberulata (Camellia, Theaceae) combined with their fourier transform infrared spectroscopy (FTIR) have been conducted in order to investigate interspecific variations which were useful in species taxonomic treatment. Based on the taxonomic treatments, chlorophyll fluorescence technology was used to study leaf chlorophyll fluorescence characteristics of seasonal changes of Tuberculata speices, and evaluate the their growth adaptation ability. FTIR was used to analyze the Phytochemical perspectives of the Tuberculata species and tea (C.sinensis) in order to evaluate their values of the application. Our results were as follows:
(1) Results of the study showed that there was considerable anatomical variation between the 18 species studied and some anatomical characters had diagnostic value in the section. The epidermal cells of Tuberculata were usually polygonal or irregular in form, with anticlinal walls straight to curved, or sinuous. The pattern of anticlinial wall varied not only in different species but also between adaxial (Ad) and abaxial (Ab) epidermis of the same species. All species in section Tuberculata were hypostomatic, and the stomata were all of the anisocytic type. All the leaves in the sections were bifacial and were composed of wax, epidermis, palisade parenchyma, spongy parenchyma, and vascular tissue. A multiple epidermis was found in most species. Sclereids were found in all 18 species studied. These results also supported Ming's merging some species in Tuberculata. All species were marked by glands and crystals, except C. ilicifolia, which supported Chang's regarded C. ilicifolia as a sole specie. Furthermore, leaf epdermic and anatomical characters were found slight difference among Tuberculata species, which indicated that Tuberculata was a natural group with close genetic relationship.
(2) Multivariate analysis (PCA and Cluster analysis) of leaf FTIR data in Tuberculata showed a visualization of the degree of affinity among the species in this section. PCA and Cluster analysis showed that C. ilicifolia became significantly distance from other 17 species. These results did not support Ming and Zhong's conclusion in transferring C. ilicifolia Y. K. Li ex H. T. Chang from the section Pseudocamellia in Chang's classification into the section Tuberculata. Multivariate analysis of leaf FTIR data in Tuberculata showed that it permitted a visualization of the degree of affinity among the species, which supported Ming's mergering of Tuberculata species.
(3) Our study indicated that the combination of characters based on leaf comparative anatomy and FTIR analysis and biogeographical distribution were useful in species revision. The results determined 11 species including C. tuberculata Chien, C. rhytidocarpa Chang et, C. rhytidophylla Chang Liang, C. anlungensis Chang, C. rubituberculata Chang, C. acutiperulata Chang et Ye., C. neriiflolia Chang, C. obovatifolia Chang, C. rubimuricata Chang et Z. R. XU, C. parvimuricata Chang, C. pyxidiacea Z. R. Xu et al. in the section Tuberculata. Moreover, our study also supported that Chang transfer C. ilicifolia Y. K. Li ex H. T. Chang from Tuberculata into the section Pseudocamellia.
(4) Leaf light-response curves of chlorophyll fluorescence, daily changes of seasonal Yield and determination of relative content of chlorophyll indicated C. rhytidophylla、C. rubituberculata、C. tuberculata、C. rhytidocarpa had a good ability to utilize light energy. Therefore, they could well adapt to grow in the region. Our studies also indicated that the ability of photosynthesis in C. neriiflolia、C. obovatifolia、C. parvimuricata were significantly inefficient, which suggested that these three species were not suitable for growth in the region. Thus, we thought that the appropriate treatment of the shade would help Tuberculata plants grown better.
(5) FTIR spectra of tea and Tuberculata species showed that carbohydrates and polysaccharides ( 3400 cm~(-1) ,1150-1030 cm~(-1)) in Tuberculata species leaves were more abundance than that in tea, indicating that vascular tissues of leaves of Tuberculata species developed well than tea. Furthermore, Tuberculata species had rich absorption peaks in 1800-1200 cm~(-1), indicating they also had rich active ingredients of tea. Therefore, they could be considered as a plant material extracted the active ingredients.
本课题通过对瘤果茶组叶的微形态、叶解剖研究,并结合傅立叶红外转换光谱(FTIR)技术,对瘤果茶组现有的存在争议的18个种进行研究,为瘤果茶组植物分类研究提供新的证据,从而澄清瘤果茶组植物分类的一些问题。同时在明确原种的基础上,进一步利用叶绿素荧光技术研究瘤果茶组植物叶片叶绿素荧光特性的季节性生长变化,对瘤果茶组种质资源从光合特性的角度进行生长适应性评估;利用FTIR方法,对瘤果茶组种质资源从植物化学的角度进行功能性的评估,最终为瘤果茶组植物资源的开发和应用提供初步的理论基础和实验证据。我们的实验结果如下:
(1)被研究的18种植物材料叶上表皮细胞形态有不规则形和多边形两种,垂周壁样式为浅波、深波和直曲型3种。下表皮细胞样式比较单一,所有种的下表皮细胞形态均为不规则形,垂周壁样式均为浅波形。所有植物的叶的气孔均分布在叶下表皮,上表皮没有气孔分布。所有气孔器类型均为环列型(anisocytic type),部分叶片具有表皮毛。所有研究的植物均为异面叶,一般由蜡质层、上下表皮、栅栏组织、海绵组织和维管组织组成。叶片栅栏组织细胞层数比较稳定。大多数种被发现有复表皮的存在。所有叶肉中均含有石细胞。除冬青叶光果茶C.ilicifolia外,其它被研究植物叶肉中均发现晶体的存在。这些结果表明叶微形态解剖特征在种的鉴定上具有一定的应用价值,可以支持闵天禄系统对部分瘤果茶植物的归并问题。同时,研究也表明叶微形态解剖特征在组间的区分上表现出更明显的分类学意义,支持张宏达系统中将冬青叶光果茶从瘤果茶组中分出的观点。此外,瘤果茶组在叶表皮微形态和叶解剖特征变化较小,也表明瘤果茶组是一个亲缘关系极为接近的、自然的组。
(2)瘤果茶组植物叶FTIR数据的PCA和聚类分析表明,冬青叶光果茶和其它17种瘤果茶有明显的差异。这一结果并不支持闵天禄将冬青叶光果茶从张宏达系统的半宿萼茶组Pseudocamellia中分离出来合并至瘤果茶组中。FTIR结果也表明,不同类群和种间的聚集程度各不相同,体现了种间的差异性和相似度,部分的支持了闵天禄系统中对瘤果茶组植物进行归并的观点。
(3)基于叶表皮微形态、叶解剖和FTIR分析的瘤果茶组种的修订结果表明:瘤果茶C.tuberculata Chien,皱果茶C.rhytidocarpa Chang et,皱叶瘤果茶C.rhytidophylla Chang Liang,安龙瘤果茶C.anlungensis Chang,厚壳红瘤果茶C.rubituberculata Chang,尖苞瘤果茶C.acutiperulata Chang et Ye.,狭叶瘤果茶C.neriiflolia Chang,倒卵叶瘤果茶C.obovatifolia Chang,荔波红瘤果茶C.rubimuricataChang et Z.R.XU,小瘤果茶C.parvimuricata Chang,三江瘤果茶C.pyxidiacea Z.R.Xu et al.等11种瘤果茶被确立。同时,研究结果也支持张宏达系统将冬青叶光果茶从瘤果茶组中移入半宿萼茶组的处理。
(4)瘤果茶组植物叶片叶绿素荧光响应曲线、季节性Yield日进程及叶绿素相对含量测定的结果表明,皱叶瘤果茶、厚壳红瘤果茶、瘤果茶、皱果茶等具有较好的光能利用能力和叶绿素生长指标,因而对于本地区的环境有较强的适应性。而狭叶瘤果茶、倒卵叶瘤果茶、小瘤果茶、尖萼瘤果茶等则表现出在本地区自然条件下生长的不适应性。因此,我们认为狭叶瘤果茶、倒卵叶瘤果茶、小瘤果茶、尖苞瘤果茶等植物类群并不适合在本地区栽种,至少不适合在野外无遮荫处理栽种。同时,在本地区栽培种植瘤果茶植物过程中,适当的遮荫处理将会有助于植物更好生长。
(5)瘤果茶组植物与茶的叶FTIR光谱显示,瘤果茶植物植物叶片红外光谱在表征碳水化合物和和多糖的3400 cm~(-1),1150-1030 cm~(-1)处的吸收强度较大,这表明瘤果茶植物叶片的维管组织比茶发达。瘤果茶植物和茶在1730 cm~(-1)附近,1650 cm~(-1),1550-1510 cm~(-1),1450和1320 cm~(-1)附近,1300-1200 cm~(-1)等光谱区域有对应的吸收峰,这些吸收峰分别表征的是茶多酚、氨基酸、茶醇、羧酸、咖啡碱等活性成分,这表明瘤果茶组植物叶也含有较为丰富的活性物质,可以考虑将其作为茶多酚、氨基酸、茶醇、咖啡碱等活性物质的来源植物。
The section Tuberculata Chang, belonging to family Theacea, genus Camellia, only distributed in China, represents a good example of a taxonomic group in the genus Camellia with controversial circumscription. This section was charactered by the tuberculate presented on the ovaries and seeds, and it was therefore considered to be orignal groups in Camellia. At present, the taxonomic treatment of the section Tuberculata is still ambiguous, which causes great inconvenience to protection of the germplasm resources and the development and utilization.
In present study, the leaf epidermial micro-morphology and comparative anatomy of 18 disputed species of the section Tuberulata (Camellia, Theaceae) combined with their fourier transform infrared spectroscopy (FTIR) have been conducted in order to investigate interspecific variations which were useful in species taxonomic treatment. Based on the taxonomic treatments, chlorophyll fluorescence technology was used to study leaf chlorophyll fluorescence characteristics of seasonal changes of Tuberculata speices, and evaluate the their growth adaptation ability. FTIR was used to analyze the Phytochemical perspectives of the Tuberculata species and tea (C.sinensis) in order to evaluate their values of the application. Our results were as follows:
(1) Results of the study showed that there was considerable anatomical variation between the 18 species studied and some anatomical characters had diagnostic value in the section. The epidermal cells of Tuberculata were usually polygonal or irregular in form, with anticlinal walls straight to curved, or sinuous. The pattern of anticlinial wall varied not only in different species but also between adaxial (Ad) and abaxial (Ab) epidermis of the same species. All species in section Tuberculata were hypostomatic, and the stomata were all of the anisocytic type. All the leaves in the sections were bifacial and were composed of wax, epidermis, palisade parenchyma, spongy parenchyma, and vascular tissue. A multiple epidermis was found in most species. Sclereids were found in all 18 species studied. These results also supported Ming's merging some species in Tuberculata. All species were marked by glands and crystals, except C. ilicifolia, which supported Chang's regarded C. ilicifolia as a sole specie. Furthermore, leaf epdermic and anatomical characters were found slight difference among Tuberculata species, which indicated that Tuberculata was a natural group with close genetic relationship.
(2) Multivariate analysis (PCA and Cluster analysis) of leaf FTIR data in Tuberculata showed a visualization of the degree of affinity among the species in this section. PCA and Cluster analysis showed that C. ilicifolia became significantly distance from other 17 species. These results did not support Ming and Zhong's conclusion in transferring C. ilicifolia Y. K. Li ex H. T. Chang from the section Pseudocamellia in Chang's classification into the section Tuberculata. Multivariate analysis of leaf FTIR data in Tuberculata showed that it permitted a visualization of the degree of affinity among the species, which supported Ming's mergering of Tuberculata species.
(3) Our study indicated that the combination of characters based on leaf comparative anatomy and FTIR analysis and biogeographical distribution were useful in species revision. The results determined 11 species including C. tuberculata Chien, C. rhytidocarpa Chang et, C. rhytidophylla Chang Liang, C. anlungensis Chang, C. rubituberculata Chang, C. acutiperulata Chang et Ye., C. neriiflolia Chang, C. obovatifolia Chang, C. rubimuricata Chang et Z. R. XU, C. parvimuricata Chang, C. pyxidiacea Z. R. Xu et al. in the section Tuberculata. Moreover, our study also supported that Chang transfer C. ilicifolia Y. K. Li ex H. T. Chang from Tuberculata into the section Pseudocamellia.
(4) Leaf light-response curves of chlorophyll fluorescence, daily changes of seasonal Yield and determination of relative content of chlorophyll indicated C. rhytidophylla、C. rubituberculata、C. tuberculata、C. rhytidocarpa had a good ability to utilize light energy. Therefore, they could well adapt to grow in the region. Our studies also indicated that the ability of photosynthesis in C. neriiflolia、C. obovatifolia、C. parvimuricata were significantly inefficient, which suggested that these three species were not suitable for growth in the region. Thus, we thought that the appropriate treatment of the shade would help Tuberculata plants grown better.
(5) FTIR spectra of tea and Tuberculata species showed that carbohydrates and polysaccharides ( 3400 cm~(-1) ,1150-1030 cm~(-1)) in Tuberculata species leaves were more abundance than that in tea, indicating that vascular tissues of leaves of Tuberculata species developed well than tea. Furthermore, Tuberculata species had rich absorption peaks in 1800-1200 cm~(-1), indicating they also had rich active ingredients of tea. Therefore, they could be considered as a plant material extracted the active ingredients.
引文
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