西藏高原硬叶柳叶片结构对寒旱环境的适应机制
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摘要
为探究硬叶柳叶片结构对高寒高海拔环境的适应机制,以西藏那曲嘉黎县硬叶柳叶片为试验材料,应用石蜡切片技术和植物显微技术,分析硬叶柳叶片结构对寒旱环境的适应特征。结果显示:(1)硬叶柳叶片为异面叶,叶片背面有表皮毛,叶片主叶脉在叶片下部凸起,呈现不规则半圆形;栅栏组织呈柱状紧密排列,有2～3层,海绵组织形状多为短柱状或者短球形,细胞间隙较大。(2)随着海拔的不断升高,硬叶柳叶片厚度从最低海拔(4 000 m)的40.4μm增加到最高海拔(4 700 m)的68.04μm,其中叶片下表皮厚度在海拔4 400 m～4 700 m之间呈显著增加的趋势,而上表皮厚度在4 000～4 400 m之间变化不显著;海绵组织和栅栏组织厚度变化分别在9.34～24.84μm、25.63～40.36μm之间,且随海拔梯度的升高海绵组织和栅栏组织的厚度均呈不显著的增加趋势。(3)硬叶柳叶片上表皮无气孔分布,下表皮散生着大量气孔,且气孔器小而密集,其密度为17.87～28.37个·mm~(-2),单位面积的气孔总周长为0.13～0.25 mm,但气孔密度和单位面积的气孔周长在海拔梯度上没有明显变化。研究表明,硬叶柳叶片主要通过增加叶片厚度来适应高海拔地区的寒旱环境。
In order to explore the adaptation mechanism of leaf structure of Salix sclerophylla to cold and high altitude environment, we used the leaf of S. sclerophylla in Jiali County, Naqu, Tibet as experimental material to study the adaptation characteristics of leaf structure of S. sclerophylla to cold and drought environment by paraffin section technology and plant microscopy technology. The results showed that:(1) the leaves of S. sclerophylla willow are heterophylous leaves with epidermis on the back of the leaves. The main veins of the leaves are convex in the lower part of the leaves, showing irregular semicircles. The palisade tissue is arranged in a column, with 2 to 3 layers. The shape of the sponge tissue is mostly short columnar or short spherical, and the cell gap is large.(2) With the increase of altitude, the leaf thickness of the S. sclerophylla willow increased from 40.4 μm at the lowest altitude(4 000 m) to 68.04 μm at the highest altitude(4 700 m). The thickness of the epidermis of the leaves increased significantly between 4 400 m and 4 700 m above sea level, but the thickness of the upper epidermis varies in not significant from 4 000 to 4400 m change. The thickness of sponge tissue and palisade tissue were between 9.34-24.84 μm and 25.63-40.36 μm, respectively. The thickness of sponge tissue and palisade tissue increased with altitude gradient, but change is not significant.(3) There is no stomatal distribution on the upper epidermis of the leaves of S. sclerophylla willow, and a large number of stomata are scattered in the lower epidermis. The stoma density is 17.87-28.37 No·mm~(-2), and the total stoma perimeter per unit area is 0.13-0.25 mm, but the stoma density and the total stoma perimeter per unit area have no significant change in the elevation gradient. Studies have shown that the leaves of S. sclerophylla willows mainly adapt to the cold and drought environment in high altitude areas by increasing the thickness of the leaves.
In order to explore the adaptation mechanism of leaf structure of Salix sclerophylla to cold and high altitude environment, we used the leaf of S. sclerophylla in Jiali County, Naqu, Tibet as experimental material to study the adaptation characteristics of leaf structure of S. sclerophylla to cold and drought environment by paraffin section technology and plant microscopy technology. The results showed that:(1) the leaves of S. sclerophylla willow are heterophylous leaves with epidermis on the back of the leaves. The main veins of the leaves are convex in the lower part of the leaves, showing irregular semicircles. The palisade tissue is arranged in a column, with 2 to 3 layers. The shape of the sponge tissue is mostly short columnar or short spherical, and the cell gap is large.(2) With the increase of altitude, the leaf thickness of the S. sclerophylla willow increased from 40.4 μm at the lowest altitude(4 000 m) to 68.04 μm at the highest altitude(4 700 m). The thickness of the epidermis of the leaves increased significantly between 4 400 m and 4 700 m above sea level, but the thickness of the upper epidermis varies in not significant from 4 000 to 4400 m change. The thickness of sponge tissue and palisade tissue were between 9.34-24.84 μm and 25.63-40.36 μm, respectively. The thickness of sponge tissue and palisade tissue increased with altitude gradient, but change is not significant.(3) There is no stomatal distribution on the upper epidermis of the leaves of S. sclerophylla willow, and a large number of stomata are scattered in the lower epidermis. The stoma density is 17.87-28.37 No·mm~(-2), and the total stoma perimeter per unit area is 0.13-0.25 mm, but the stoma density and the total stoma perimeter per unit area have no significant change in the elevation gradient. Studies have shown that the leaves of S. sclerophylla willows mainly adapt to the cold and drought environment in high altitude areas by increasing the thickness of the leaves.
引文
[1] 李全发,王宝娟,安丽华,等.青藏高原草地植物叶解剖特征[J].生态学报,2013,33(7):2 062-2 070.LI Q F,WANG B J,AN L H,et al.Leaf anatomical characteristics of the plants of grasslands in the Tibetan Plateau[J].Acta Ecologica Sinica,2013,33(7):2 062-2 070.
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[3] 李芳兰,包维楷.植物叶片形态解剖结构对环境变化的响应与适应[J].植物学通报,2005(B08):118-127.LI F L,BAO W K.Responses of the morphological and anatomical structure of the plant leaf to environmental change[J].Chinese Bulletin of Botany,2005,(B08):118-127.
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[5] 姜永雷,邓莉兰,黄晓霞.不同海拔川滇高山栎叶片的解剖结构特征[J].江苏农业科学,2015,43(1):195-198.JIANG Y L,DENG L L,HUANG X X.Leaf anatomical characteristics of the plants of Quercus aquifolioides in the different altitudes[J].Jiangsu Agricultural Sciences,2015,43(1):195-198.
[6] 刘培卫,张玉秀,杨云,等.六种沉香属植物叶片解剖结构研究[J].广西植物,2017,37(5):565-571.LIU P W,ZHANG Y X,YANG Y,et al.Leaf anatomical structure of six Aquilaria species[J].Guihaia,2017,37(5):565-571.
[7] 朱广龙,魏学智.酸枣叶片结构可塑性对自然梯度干旱生境的适应特征[J].生态学报,2016,36(19):6 178-6 187.ZHU G L,WEI X Z.Leaf morphological plasticity of Ziziphus jujuba var.spinosa in response to natural drought gradient ecotopes[J].Acta Ecologica Sinica,2016,36(19):6 178-6 187.
[8] 朴世龙.青藏高原草地植被覆盖变化及其与气候因子的关系[J].植物生态学报,2006,30(1):1-8.PIAO S L.Variations in grassland vegetation cover in relation to climatic factors on the Tibetan plateau[J].Journal of Plant Ecology,2006,30(1):1-8.
[9] 李文华,周兴民.青藏高原生态系统及优化利用模式[M].广州:广东科技出版社,1998.
[10] KLICH M G.Leaf variations in Elaeagnus angustifolia related to environmental heterogeneity[J].Environmental and Experimental Botany,2000,44(3):171-183.
[11] 王瑞丽,于贵瑞,何念鹏,等.气孔特征与叶片功能性状之间关联性沿海拔梯度的变化规律:以长白山为例[J].生态学报,2016,36(8):2 175-2 184.WANG R L,YU G R,HE N P,et al.Altitudinal variation in the covariation of stomatal traits with leaf functional traits in Changbai Mountain[J].Acta Ecologica Sinica,2016,36(8):2 175-2 184.
[12] 张秀芳,石东里,张兰.观察植物气孔结构的简易方法[J].生物学通报,2002,37(6):42.ZHANG X F,SHI D L,ZHANG L.Simple method of observing stomatal structure [J].Bulletin of Biology,2002,37(6):42.
[13] 宋丽清,胡春梅,侯喜林,等.高梁、紫苏叶脉密度与光合特性的关系[J].植物学报,2015,50(1):100-106.SONG L Q,HU C M,HOU X L,et al.Relationship between photosynthetic characteristics and leaf vein density in Sorghum bicolor and Perilla frutescens[J].Bulletin of Botany,2015,50(1):100-106.
[14] BEAULIEU J M,LEITCH I J,PATEL S,et al.Genome size is a strong predictor of cell size and stomatal density in angiosperms[J].The New Phytologist,2008,179(4):975-986.
[15] 李国华,徐涛,陈国云,等.10个品种澳洲坚果叶片解剖学的比较研究[J].热带作物学报,2009,30(10):1 437-1 441.LI G H,XU T,CHEN G Y,et al.Anatomical structure of leaves of 10 macadamia cultivars[J].Chinese Journal of Tropical Crops,2009,30(10):1 437-1 441.
[16] 卡特.植物解剖学(下册)[M].李正理译,北京:科学出版社,1976.
[17] 王勋陵,王静.植物形态结构与环境[M].兰州:兰州大学出版社,1989:205-230.
[18] 沈广爽,石雪芹,古松,等.九种海滨沙生植物解剖构造及其生态适应性研究[J].广西植物,2014,34(2):263-268.SHEN G S,SHI X Q,GU S,et al.Anatomical structures and ecological adaptability research of nine species of psammophytes on beach[J].Guihaia,2014,34(2):263-268.
[19] 李芳兰,包维楷,刘俊华.岷江上游干旱河谷海拔梯度上四川黄栌叶片特征及其与环境因子的关系[J].西北植物学报,2005,25(11):2 277-2 284.LI F L,BAO W K,LIU J H.Leaf characteristics and their relationship of Cotinus coggygria in arid river valley located in the upper reaches of Minjiang River with environmental factors depending on its altitude gradients[J].Acta Botanica Boreali-Occidentalia Sinica,2005,25(11):2 277-2 284.
[20] 王国严,徐阿生.川滇高山栎研究综述[J].四川林业科技,2008,29(2):23-29,34.WANG G Y,XU A S.A review of researches on Quercus aquifolioides[J].Journal of Sichuan Forestry Science and Technology,2008,29(2):23-29,34.
[21] 贺金生,陈伟烈,王勋陵.高山栎叶的形态结构及其与生态环境的关系[J].植物生态学报,1994,18(3):219-227.HE J S,CHEN W L,WANG X L.Morphological and anatomical features of Quercus section suber and its adaptation to the ecological environment[J].Acta Phytoecologica Sinica,1994,18(3):219-227.
[22] 容丽,王世杰,刘宁,等.喀斯特山区先锋植物叶片解剖特征及其生态适应性评价:以贵州花江峡谷区为例[J].山地学报,2005,23(1):35-42.RONG L,WANG S J,LIU N,et al.Leaf anatomical characters and its ecological adaptation of the pioneer species in the karst mountain area:with a special reference to the huajiang canyon of Guizhou[J].Journal of Mountain Science,2005,23(1):35-42.
[23] CHEN Y N,WANG Q,RUAN X,et al.Physiological Response of Populus euphratica to artificial water-recharge of the lower reaches of Tarim River[J].Acta Botanica Sinica,2004,46(12):1 393-1 401.
[24] SAM O,JERZ E,DELLAM ICO J,et al.Water stress induced changes in anatomy of tomato leaf epidermis[J].Biologia Plantarum,2000,(43):275-277.
[25] 费松林,方精云,樊拥军,等.贵州梵净山亮叶水青冈叶片和木材的解剖学特征及其与生态因子的关系[J].植物学报,1999,41(9):1 002-1 009.FEI S L,FANG J Y,FAN Y J,et al.Anatomical characteristics of leaves and woods of Fagus lucida and their relationship to ecological factors in Mountain Fanjingshan,Guizhou,China[J].Acta Botanica Sinica,1999,41(9):1 002-1 009.
[2] MOTT K A,GIBSON A C,OLEARY J W.The adaptive significance of amphistomatic leaves[J].Plant Cell and Environment,1982,5(6):445-460.
[3] 李芳兰,包维楷.植物叶片形态解剖结构对环境变化的响应与适应[J].植物学通报,2005(B08):118-127.LI F L,BAO W K.Responses of the morphological and anatomical structure of the plant leaf to environmental change[J].Chinese Bulletin of Botany,2005,(B08):118-127.
[4] 郭学民,刘建珍,翟江涛,等.16个品种桃叶片解剖结构与树干抗寒性的关系[J].林业科学,2015,51(8):33-43.GUO X M,LIU J Z,ZHAI J T,et al.Relationship between leaf anatomical structure and trunk cold resistance of 16 peach cultivars[J].Scientia Silvae Sinicae,2015,51(8):33-43.
[5] 姜永雷,邓莉兰,黄晓霞.不同海拔川滇高山栎叶片的解剖结构特征[J].江苏农业科学,2015,43(1):195-198.JIANG Y L,DENG L L,HUANG X X.Leaf anatomical characteristics of the plants of Quercus aquifolioides in the different altitudes[J].Jiangsu Agricultural Sciences,2015,43(1):195-198.
[6] 刘培卫,张玉秀,杨云,等.六种沉香属植物叶片解剖结构研究[J].广西植物,2017,37(5):565-571.LIU P W,ZHANG Y X,YANG Y,et al.Leaf anatomical structure of six Aquilaria species[J].Guihaia,2017,37(5):565-571.
[7] 朱广龙,魏学智.酸枣叶片结构可塑性对自然梯度干旱生境的适应特征[J].生态学报,2016,36(19):6 178-6 187.ZHU G L,WEI X Z.Leaf morphological plasticity of Ziziphus jujuba var.spinosa in response to natural drought gradient ecotopes[J].Acta Ecologica Sinica,2016,36(19):6 178-6 187.
[8] 朴世龙.青藏高原草地植被覆盖变化及其与气候因子的关系[J].植物生态学报,2006,30(1):1-8.PIAO S L.Variations in grassland vegetation cover in relation to climatic factors on the Tibetan plateau[J].Journal of Plant Ecology,2006,30(1):1-8.
[9] 李文华,周兴民.青藏高原生态系统及优化利用模式[M].广州:广东科技出版社,1998.
[10] KLICH M G.Leaf variations in Elaeagnus angustifolia related to environmental heterogeneity[J].Environmental and Experimental Botany,2000,44(3):171-183.
[11] 王瑞丽,于贵瑞,何念鹏,等.气孔特征与叶片功能性状之间关联性沿海拔梯度的变化规律:以长白山为例[J].生态学报,2016,36(8):2 175-2 184.WANG R L,YU G R,HE N P,et al.Altitudinal variation in the covariation of stomatal traits with leaf functional traits in Changbai Mountain[J].Acta Ecologica Sinica,2016,36(8):2 175-2 184.
[12] 张秀芳,石东里,张兰.观察植物气孔结构的简易方法[J].生物学通报,2002,37(6):42.ZHANG X F,SHI D L,ZHANG L.Simple method of observing stomatal structure [J].Bulletin of Biology,2002,37(6):42.
[13] 宋丽清,胡春梅,侯喜林,等.高梁、紫苏叶脉密度与光合特性的关系[J].植物学报,2015,50(1):100-106.SONG L Q,HU C M,HOU X L,et al.Relationship between photosynthetic characteristics and leaf vein density in Sorghum bicolor and Perilla frutescens[J].Bulletin of Botany,2015,50(1):100-106.
[14] BEAULIEU J M,LEITCH I J,PATEL S,et al.Genome size is a strong predictor of cell size and stomatal density in angiosperms[J].The New Phytologist,2008,179(4):975-986.
[15] 李国华,徐涛,陈国云,等.10个品种澳洲坚果叶片解剖学的比较研究[J].热带作物学报,2009,30(10):1 437-1 441.LI G H,XU T,CHEN G Y,et al.Anatomical structure of leaves of 10 macadamia cultivars[J].Chinese Journal of Tropical Crops,2009,30(10):1 437-1 441.
[16] 卡特.植物解剖学(下册)[M].李正理译,北京:科学出版社,1976.
[17] 王勋陵,王静.植物形态结构与环境[M].兰州:兰州大学出版社,1989:205-230.
[18] 沈广爽,石雪芹,古松,等.九种海滨沙生植物解剖构造及其生态适应性研究[J].广西植物,2014,34(2):263-268.SHEN G S,SHI X Q,GU S,et al.Anatomical structures and ecological adaptability research of nine species of psammophytes on beach[J].Guihaia,2014,34(2):263-268.
[19] 李芳兰,包维楷,刘俊华.岷江上游干旱河谷海拔梯度上四川黄栌叶片特征及其与环境因子的关系[J].西北植物学报,2005,25(11):2 277-2 284.LI F L,BAO W K,LIU J H.Leaf characteristics and their relationship of Cotinus coggygria in arid river valley located in the upper reaches of Minjiang River with environmental factors depending on its altitude gradients[J].Acta Botanica Boreali-Occidentalia Sinica,2005,25(11):2 277-2 284.
[20] 王国严,徐阿生.川滇高山栎研究综述[J].四川林业科技,2008,29(2):23-29,34.WANG G Y,XU A S.A review of researches on Quercus aquifolioides[J].Journal of Sichuan Forestry Science and Technology,2008,29(2):23-29,34.
[21] 贺金生,陈伟烈,王勋陵.高山栎叶的形态结构及其与生态环境的关系[J].植物生态学报,1994,18(3):219-227.HE J S,CHEN W L,WANG X L.Morphological and anatomical features of Quercus section suber and its adaptation to the ecological environment[J].Acta Phytoecologica Sinica,1994,18(3):219-227.
[22] 容丽,王世杰,刘宁,等.喀斯特山区先锋植物叶片解剖特征及其生态适应性评价:以贵州花江峡谷区为例[J].山地学报,2005,23(1):35-42.RONG L,WANG S J,LIU N,et al.Leaf anatomical characters and its ecological adaptation of the pioneer species in the karst mountain area:with a special reference to the huajiang canyon of Guizhou[J].Journal of Mountain Science,2005,23(1):35-42.
[23] CHEN Y N,WANG Q,RUAN X,et al.Physiological Response of Populus euphratica to artificial water-recharge of the lower reaches of Tarim River[J].Acta Botanica Sinica,2004,46(12):1 393-1 401.
[24] SAM O,JERZ E,DELLAM ICO J,et al.Water stress induced changes in anatomy of tomato leaf epidermis[J].Biologia Plantarum,2000,(43):275-277.
[25] 费松林,方精云,樊拥军,等.贵州梵净山亮叶水青冈叶片和木材的解剖学特征及其与生态因子的关系[J].植物学报,1999,41(9):1 002-1 009.FEI S L,FANG J Y,FAN Y J,et al.Anatomical characteristics of leaves and woods of Fagus lucida and their relationship to ecological factors in Mountain Fanjingshan,Guizhou,China[J].Acta Botanica Sinica,1999,41(9):1 002-1 009.