杉科植物的表皮结构及其气孔参数分析
摘要
本文主要研究杉科植物的表皮结构,建立杉科化石植物分类鉴定的表皮结构参照系;分析红杉属、水杉属、水松属和落羽杉属植物的气孔参数,根据统计学的结果定量分析在指示大气CO_2浓度的研究工作中,气孔密度和气孔指数哪个更可靠;利用标本的枝叶和表皮结构的特征,确定采自中国黑龙江依兰始新世和云南吕合中新世地层杉科化石植物标本的分类地位。
杉科常绿植物的叶片通常为革质,落叶植物的叶片为纸质,落叶植物的叶片角质层比常绿植物的叶片角质层薄。非气孔区的表皮细胞排成规则的纵向行列,细胞长方形,细胞长轴与叶片长轴一致。水杉属的多数表皮细胞垂周壁明显弯曲,落羽杉属和杉木属有时微呈波状,其他属的表皮细胞垂周壁直。气孔区的表皮细胞形状有时为多边形,细胞长轴多数也与叶片长轴平行。叶片为双面气孔型或单面气孔型,多数植物的叶片近轴面和远轴面的气孔数量和分布不同。一般来说,条形叶和披针形叶的远轴面气孔分布状况和气孔数量稳定,远轴面的中部最稳定。条形叶远轴面的气孔分布于中脉两侧,形成纵向的气孔带。条形叶近轴面气孔分布状况和气孔数量变化大,近轴面的气孔数量有时与远轴面近似,但多数情况下比远轴面少,有时整个叶片的近轴面仅少数几个气孔或没有气孔分布。钻形叶的近轴面和远轴面的气孔数量近似,或叶片近轴面的气孔数量比远轴面的气孔数量多,气孔分布范围也比远轴面气孔分布范围广。气孔椭圆形,落羽杉属植物的气孔长轴方向与叶片长轴垂直或斜向排列,柳杉属植物的气孔多斜向排列。落羽杉属和柳杉属以外的杉科植物的气孔长轴多数与叶片长轴平行。保卫细胞下陷,细胞垂周壁加厚明显。保卫细胞的极端联合,细胞垂周壁向极部延长并向2侧弯曲。气孔的副卫细胞通常为4-7个。
根据对广州和杭州2个采集地水松标本气孔参数的t-检验结果,条形叶的气孔参数的变化比条状钻形叶气孔参数的变化小。条状钻形叶远轴面气孔指数的变化比近轴面气孔指数的变化小,也比近轴面和远轴面气孔密度的变化小。因此在指示大气CO_2浓度的工作中,用水松的条形叶比条状钻形叶可靠,用气孔指数比用气孔密度可靠,用条状钻形叶的远轴面气孔指数比近轴面的气孔指数可靠。许多植物的叶内不同部位气孔参数也不同,分析从相同部位获取的北美红杉、水杉、水松和落羽杉的气孔参数发现,气孔指数的变异系数小于气孔密度的变异系数,气孔指数的稳定性比气孔密度的稳定性高。不论是定性分析还是t-检验和变异系数分析等定量研究的结果都表明,在利用气孔参数指示大气CO_2浓度的工作中,用气孔指数指示大气CO_2浓度比用气孔密度指示更为准确。
中国黑龙江依兰始新世和云南吕合中新世地层发现的标本的叶片形态和排列与杉科的红杉属、水杉属、水松属和落羽杉属植物相似。水松属和落羽杉属植物条形叶的质地为纸质,叶尖比较尖锐,基部下延部分不明显,平行沿小枝轴延伸等特征
The objective of the present work is to establish a frame of reference in epidermal structures of Taxodiaceae, and to determine quantitatively the reliable stomatal parameter (density or index) as a CO_2 indicator in genera Sequoia, Metasequoa, Taxodium and Glyptostrobus based on statistical analysis, and to reveal the affinity of speciemen from the Eocene of Yilan, Heilongjiang Province, NE China and from the Miocene of Luhe, Yunnan Province, South China.The deciduous genera in Taxodiaceae commonly have papyraceous cuticle which is easily distinguished from coriaceous cuticle of evergreen leaves. Epidermal cells of stomata-free area are oblong and parallel to the long axis of leaves. The anticlinal walls of epidermal cells are mostly straight in most genera, sometimes slightly undulate in Taxodium and Cunninghamia, and obviously undulate in Metasequoia. Epidermal cells on stomatal areas are sometimes polygonous. Pits can be clearly seen in the walls of epidermal cells. Leaves are amphistomatic or hypostomatic. Stomata are located on each side of the mid-vein. The numbers of stomata on the abaxial surfaces are generally more than or sometimes the same as those on the adaxial surfaces in linear and lanceolate leaves. The distribution of stomata is generally more uniform on the abaxial surface of linear and lanceolate leaves, and the mid-part of the abaxial surface yields the least variation in stomatal distribution. The numbers of stomata on the adaxial surfaces are more than or sometimes the same as those on the abaxial surfaces in subulate leaves of Taxodiaceae. Stomatal apparatus are oval, and the long axes of stomata are perpendicular or oblique to the midvein of the leaves in Taxodium, and oblique to the midvein of the leaves in Cryptomeria, mostly parallel to the midvein of the leaves in other genera of Taxodiaceae. Guard cells have thickened walls, especially on outer margins. The thickened walls of guard cells form polar lamellae that produce protruding and curved ends at the two poles of the stomata. Commonly the stomata have 4 to 7 subsidiary cells.The Mest of stomatal parameters of Glyptostrobus pensilis from Guangzhou and Hangzhou indicate the variation of stomatal parameters of linear leaves is smaller than that of linear-subulate leaves. The variation of stomatal index of the abaxial surfaces is smaller than that of the index of the adaxial surface, and also is smaller than that of the density of the abaxial and adaxial surfaces in linear-subulate leaves according to the Mest. When studying the correlation between stomatal parameters of Glyptostrobus and atmospheric CO_2 concentrations, the stomatal parameters of linear leaves are firstly chosen, and the stomatal index is more useful than stomatal density. The intra-individual variation of stomatal index is smaller than that of the density by comparing the coefficient of variability
(CV) of stomatal density with the CV of index in genera Sequoia, Metasequoa, Taxodium and Glyptostrobus. Thus, even if stomatal density and stomatal index show similar responses for a given species, stomatal index should yield more accurate CO_2 estimates in genera of Sequoia, Taxodium, Metasequoia and Glyptostrobus.Foliage shoots of Sequoia are reported form the Eocene of Yilan, Heilongjiang Province, NE China and from the Miocene of Liihe, Yunnan Province, South China. The leaves from both Yilan and Liihe are linear in shape, coriaceous in texture and the leaf base is decurrent and attached obliquely to the axis of shoots. The walls of epidermal cells are straight, and the long axis of stomata is mostly parallel to mid-vein in present specimens. It suggests that the new specimens are assigned to Sequoia rather than to similar genera of Metasequoia, Taxodium and Glyptostrobus. The presence of Sequoia during the Eocene in Yilan and Miocene in Luhe shows the detailed cuticle, which may provides valuable information on cellular structures and on the palaeoclimate.
杉科常绿植物的叶片通常为革质,落叶植物的叶片为纸质,落叶植物的叶片角质层比常绿植物的叶片角质层薄。非气孔区的表皮细胞排成规则的纵向行列,细胞长方形,细胞长轴与叶片长轴一致。水杉属的多数表皮细胞垂周壁明显弯曲,落羽杉属和杉木属有时微呈波状,其他属的表皮细胞垂周壁直。气孔区的表皮细胞形状有时为多边形,细胞长轴多数也与叶片长轴平行。叶片为双面气孔型或单面气孔型,多数植物的叶片近轴面和远轴面的气孔数量和分布不同。一般来说,条形叶和披针形叶的远轴面气孔分布状况和气孔数量稳定,远轴面的中部最稳定。条形叶远轴面的气孔分布于中脉两侧,形成纵向的气孔带。条形叶近轴面气孔分布状况和气孔数量变化大,近轴面的气孔数量有时与远轴面近似,但多数情况下比远轴面少,有时整个叶片的近轴面仅少数几个气孔或没有气孔分布。钻形叶的近轴面和远轴面的气孔数量近似,或叶片近轴面的气孔数量比远轴面的气孔数量多,气孔分布范围也比远轴面气孔分布范围广。气孔椭圆形,落羽杉属植物的气孔长轴方向与叶片长轴垂直或斜向排列,柳杉属植物的气孔多斜向排列。落羽杉属和柳杉属以外的杉科植物的气孔长轴多数与叶片长轴平行。保卫细胞下陷,细胞垂周壁加厚明显。保卫细胞的极端联合,细胞垂周壁向极部延长并向2侧弯曲。气孔的副卫细胞通常为4-7个。
根据对广州和杭州2个采集地水松标本气孔参数的t-检验结果,条形叶的气孔参数的变化比条状钻形叶气孔参数的变化小。条状钻形叶远轴面气孔指数的变化比近轴面气孔指数的变化小,也比近轴面和远轴面气孔密度的变化小。因此在指示大气CO_2浓度的工作中,用水松的条形叶比条状钻形叶可靠,用气孔指数比用气孔密度可靠,用条状钻形叶的远轴面气孔指数比近轴面的气孔指数可靠。许多植物的叶内不同部位气孔参数也不同,分析从相同部位获取的北美红杉、水杉、水松和落羽杉的气孔参数发现,气孔指数的变异系数小于气孔密度的变异系数,气孔指数的稳定性比气孔密度的稳定性高。不论是定性分析还是t-检验和变异系数分析等定量研究的结果都表明,在利用气孔参数指示大气CO_2浓度的工作中,用气孔指数指示大气CO_2浓度比用气孔密度指示更为准确。
中国黑龙江依兰始新世和云南吕合中新世地层发现的标本的叶片形态和排列与杉科的红杉属、水杉属、水松属和落羽杉属植物相似。水松属和落羽杉属植物条形叶的质地为纸质,叶尖比较尖锐,基部下延部分不明显,平行沿小枝轴延伸等特征
The objective of the present work is to establish a frame of reference in epidermal structures of Taxodiaceae, and to determine quantitatively the reliable stomatal parameter (density or index) as a CO_2 indicator in genera Sequoia, Metasequoa, Taxodium and Glyptostrobus based on statistical analysis, and to reveal the affinity of speciemen from the Eocene of Yilan, Heilongjiang Province, NE China and from the Miocene of Luhe, Yunnan Province, South China.The deciduous genera in Taxodiaceae commonly have papyraceous cuticle which is easily distinguished from coriaceous cuticle of evergreen leaves. Epidermal cells of stomata-free area are oblong and parallel to the long axis of leaves. The anticlinal walls of epidermal cells are mostly straight in most genera, sometimes slightly undulate in Taxodium and Cunninghamia, and obviously undulate in Metasequoia. Epidermal cells on stomatal areas are sometimes polygonous. Pits can be clearly seen in the walls of epidermal cells. Leaves are amphistomatic or hypostomatic. Stomata are located on each side of the mid-vein. The numbers of stomata on the abaxial surfaces are generally more than or sometimes the same as those on the adaxial surfaces in linear and lanceolate leaves. The distribution of stomata is generally more uniform on the abaxial surface of linear and lanceolate leaves, and the mid-part of the abaxial surface yields the least variation in stomatal distribution. The numbers of stomata on the adaxial surfaces are more than or sometimes the same as those on the abaxial surfaces in subulate leaves of Taxodiaceae. Stomatal apparatus are oval, and the long axes of stomata are perpendicular or oblique to the midvein of the leaves in Taxodium, and oblique to the midvein of the leaves in Cryptomeria, mostly parallel to the midvein of the leaves in other genera of Taxodiaceae. Guard cells have thickened walls, especially on outer margins. The thickened walls of guard cells form polar lamellae that produce protruding and curved ends at the two poles of the stomata. Commonly the stomata have 4 to 7 subsidiary cells.The Mest of stomatal parameters of Glyptostrobus pensilis from Guangzhou and Hangzhou indicate the variation of stomatal parameters of linear leaves is smaller than that of linear-subulate leaves. The variation of stomatal index of the abaxial surfaces is smaller than that of the index of the adaxial surface, and also is smaller than that of the density of the abaxial and adaxial surfaces in linear-subulate leaves according to the Mest. When studying the correlation between stomatal parameters of Glyptostrobus and atmospheric CO_2 concentrations, the stomatal parameters of linear leaves are firstly chosen, and the stomatal index is more useful than stomatal density. The intra-individual variation of stomatal index is smaller than that of the density by comparing the coefficient of variability
(CV) of stomatal density with the CV of index in genera Sequoia, Metasequoa, Taxodium and Glyptostrobus. Thus, even if stomatal density and stomatal index show similar responses for a given species, stomatal index should yield more accurate CO_2 estimates in genera of Sequoia, Taxodium, Metasequoia and Glyptostrobus.Foliage shoots of Sequoia are reported form the Eocene of Yilan, Heilongjiang Province, NE China and from the Miocene of Liihe, Yunnan Province, South China. The leaves from both Yilan and Liihe are linear in shape, coriaceous in texture and the leaf base is decurrent and attached obliquely to the axis of shoots. The walls of epidermal cells are straight, and the long axis of stomata is mostly parallel to mid-vein in present specimens. It suggests that the new specimens are assigned to Sequoia rather than to similar genera of Metasequoia, Taxodium and Glyptostrobus. The presence of Sequoia during the Eocene in Yilan and Miocene in Luhe shows the detailed cuticle, which may provides valuable information on cellular structures and on the palaeoclimate.
引文
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