黄土丘陵沟壑区10种单子叶植物叶片解剖结构及环境适应性
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摘要
该研究选取黄土丘陵沟壑区常见的10种单子叶植物,采用常规石蜡切片法,对不同植被带(草原带、森林草原带、森林带)及不同立地环境(阳坡、峁顶、阴坡)植物叶片解剖结构进行观察,并测量叶片上角质层厚度、下角质层厚度、上表皮厚度、下表皮厚度、叶片厚度、导管直径等指标,以探讨该区不同单子叶植物解剖结构特征及环境适应性。结果表明:(1)单子叶植物叶片上、下表皮外侧均有不同程度的角质层分化;叶片上表皮还有泡状细胞的分化,但不同物种泡状细胞的位置和大小不同;单子叶植物较大的维管束中一般有2～4个明显粗大的导管,呈V形排列。单子叶植物可通过自身特殊的解剖结构特征适应研究区特有的环境。(2)叶片解剖结构中仅有导管直径在不同植被带间差异显著,随土壤水分的减少,植物主要通过增大导管直径来适应不良的水分条件。(3)叶片解剖结构中表皮和上角质层厚度等保护组织在不同立地环境间有显著差异,在水分条件差、光照强烈的环境中,植物主要通过增大表皮和上角质层厚度来适应环境。
In this study,10 monocotyledons species were chosen in the hilly and gully region of the Loess Plateau.Conventional paraffin section method was used.The leaf anatomical structures of ten species in different vegetation zones(steppe zone,forest steppe zone and forest zone)and different erosion environments(south face slope,hilltop and north face slope)were observed.We measured upper cuticle thickness,upper epidermis thickness,lower epidermis thickness,lower cuticle thickness,leaf thickness,vessel diameter,and discussed anatomical structure characteristics and environmental adaptability of these species.The research results showed that:(1)there were different degrees of cuticle differentiation on the upper and lower epidermis in the ten monocotyledon plants.The leaves also had differentiation of bulliform cells in their upper epidermis.The locations and sizes of bulliform cells were different among different species.There were two or four big vessels in monocotyledon plants,which arranged in"V"-shaped in the big vascular bundles.Monocotyledon plants could adapt to the special environment in this area through their own special anatomical structure characteristics.(2)In leaf anatomical structures,only vessels diameter had significant difference among different vegetation zones.With the decrease of soil moisture,plants adapted to poor water conditions mainly by increasing vessels diameter.(3)Protective tissues,such as epidermis thickness and upper cuticle thickness in leaf anatomical structures,were significant differences in different erosion environments.Plants adapted to poor water and strong light conditions mainly by increasing the thickness of epidermis and upper cuticle.
In this study,10 monocotyledons species were chosen in the hilly and gully region of the Loess Plateau.Conventional paraffin section method was used.The leaf anatomical structures of ten species in different vegetation zones(steppe zone,forest steppe zone and forest zone)and different erosion environments(south face slope,hilltop and north face slope)were observed.We measured upper cuticle thickness,upper epidermis thickness,lower epidermis thickness,lower cuticle thickness,leaf thickness,vessel diameter,and discussed anatomical structure characteristics and environmental adaptability of these species.The research results showed that:(1)there were different degrees of cuticle differentiation on the upper and lower epidermis in the ten monocotyledon plants.The leaves also had differentiation of bulliform cells in their upper epidermis.The locations and sizes of bulliform cells were different among different species.There were two or four big vessels in monocotyledon plants,which arranged in"V"-shaped in the big vascular bundles.Monocotyledon plants could adapt to the special environment in this area through their own special anatomical structure characteristics.(2)In leaf anatomical structures,only vessels diameter had significant difference among different vegetation zones.With the decrease of soil moisture,plants adapted to poor water conditions mainly by increasing vessels diameter.(3)Protective tissues,such as epidermis thickness and upper cuticle thickness in leaf anatomical structures,were significant differences in different erosion environments.Plants adapted to poor water and strong light conditions mainly by increasing the thickness of epidermis and upper cuticle.
引文
[1]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.
[2]ACKERLY D,KNIGHT C,WEISS S,et al.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:Contrasting patterns in species level and community level analyses[J].Oecologia,2002,130(3):449-457.
[3]YIN Q L,WANG L,LEI M L,et al.The relationships between leaf economics and hydraulic traits of woody plants depend on water availability[J].The Science of the Total Environment,2018,621:245-252.
[4]BUCKLEY T N,JOHN G P,SCOFFONI C,et al.How does leaf anatomy influence water transport outside the xylem[J].Plant Physiology,2015,168(4):1 616-1 635.
[5]TOMS M,FLEXAS J,et al.Importance of leaf anatomy in determining mesophyll diffusion conductance to CO2 across species:Quantitative limitations and scaling up by models[J].Journal of Experimental Botany,2013,64(8):2 269-2 281.
[6]XIONG D L,FLEXAS J,YU T T,et al.Leaf anatomy mediates coordination of leaf hydraulic conductance and mesophyll conductance to CO2in Oryza[J].The New Phytologist,2017,213(2):572-583.
[7]陈庆诚,孙仰文,张国樑.疏勒河中、下游植物群落优势种生态-形态、解剖特性的初步研究[J].兰州大学学报,1961,(3):65-100.CHEN Q C,SUN Y W,ZHANG G L.A preliminary study on the morphology and anatomy of the dominants of the vegetations distributing along the middle and low drainage basin of the Shor-Ler River based on the view point of ecology[J].Journal of Lanzhou University,1961,(3):65-100.
[8]HE N P,LIU C C,TIAN M,et al.Variation in leaf anatomical traits from tropical to cold-temperate forests and linkage to ecosystem functions[J].Functional Ecology,2018,32(1):10-19.
[9]陈莹婷,许振柱.植物叶经济谱的研究进展[J].植物生态学报,2014,38(10):1 135-1 153.CHEN Y T,XU Z Z.Review on research of leaf economics spectrum[J].Chinese Journal of Plant Ecology,2014,38(10):1 135-1 153.
[10]FU B J,LIU Y,LY H,et al.Assessing the soil erosion control service of ecosystems change in the Loess Plateau of China[J].Ecological Complexity,2011,8(4):284-293.
[11]杜华栋.黄土丘陵沟壑区优势植物对不同侵蚀环境的适应研究---从群落、个体、组织到生理[D].北京:中国科学院大学,2013.
[12]杜华栋,焦菊英,寇萌,等.黄土高原先锋种猪毛蒿叶片形态解剖与生理特征对立地的适应性[J].生态学报,2016,36(10):2 914-2 925.DU H D,JIAO J Y,KOU M,et al.Adaptability of foliar morphological,anatomical,and physiological characteristics of the pioneer species Artemisia scoparia growing in a hillygully Loess Region at different slope sites[J].Acta Ecologica Sinica,2016,36(10):2 914-2 925.
[13]杜华栋,徐翠红,刘萍,等.陕北黄土高原优势植物叶片解剖结构的生态适应性[J].西北植物学报,2010,30(2):293-300.DU H D,XU C H,LIU P,et al.Foliar anatomical structures and ecological adaptabilities of dominant plants in the north Shaanxi Loess Plateau[J].Acta Botanica Boreali-Occidentalia Sinica,2010,30(2):293-300.
[14]王勇,梁宗锁,龚春梅,等.干旱胁迫对黄土高原4种蒿属植物叶形态解剖学特征的影响[J].生态学报,2014,34(16):4 535-4 548.WANG Y,LIANG Z S,GONG C M,et al.Effect of drought on leaf anatomical characteristics of four Artemisia species in the Loess Plateau[J].Acta Ecologica Sinica,2014,34(16):4 535-4 548.
[15]杨超,梁宗锁.陕北撂荒地上优势蒿类叶片解剖结构及其生态适应性[J].生态学报,2008,28(10):4 732-4 738.YANG C,LIANG Z S.Foliar anatomical structures and ecological adaptabilities of dominant artemisia species of early sere of succession on arable old land after being abandoned in Loess Hilly Region[J].Acta Ecologica Sinica,2008,28(10):4 732-4 738.
[16]KOU M,JIAO J Y,YIN Q L,et al.Successional trajectory over 10years of vegetation restoration of abandoned slope croplands in the hill-gully region of the Loess Plateau[J].Land Degradation&Development,2016,27(4):919-932.
[17]中国科学院黄土高原综合科学考察队.黄土高原地区土壤侵蚀区域特征及其治理途径[M].北京:中国科学技术出版社,1991.
[18]王义风.黄土高原地区植被资源及其合理利用[M].北京:科学技术出版社,1991.
[19]李和平.植物显微技术[M].2版.北京:科学出版社,2009.
[20]龚春梅,宁蓬勃,王根轩,等.C3和C4植物光合途径的适应性变化和进化[J].植物生态学报,2009,33(1):206-221.GONG C M,NING P B,WANG G X,et al.A review of adaptable variations and evolution of photosynthetic carbon assimilating pathway in C3and C4plants[J].Chinese Journal of Plant Ecology,2009,33(1):206-221.
[21]李正理.旱生植物的形态和结构[J].生物学通报,1981,16(4):9-12.LI Z L.The structure and morphology of xeromorphic[J].Bulletin of Biology,1981,16(4):9-12.
[22]黄振英,吴鸿,胡正海.30种新疆沙生植物的结构及其对沙漠环境的适应[J].植物生态学报,1997,21(6):521-530.HUANG Z Y,WU H,HU Z H.The structures of 30species of psammophytes and their adaptation to the sandy desert environment in Xinjiang[J].Acta Phytoecologica Sinica,1997,21(6):521-530.
[23]穆兴民.试论黄土区旱地土壤水资源的地带性与非地带性[J].土壤学报,1999,36(2):237-244.MU X M.An exploratory research on the geo-zonal and unzonal distributions of soil water in loess region,China[J].Acta Pedologica Sinica,1999,36(2):237-244.
[24]王力,邵明安,侯庆春.土壤干层量化指标初探[J].水土保持学报,2000,14(4):87-90.WANG L,SHAO M A,HOU Q C.Preliminary research on measured indexes of dried soil layer[J].Journal of Soil and Water Conservation,2000,14(4):87-90.
[25]马建静,吉成均,韩梅,等.青藏高原高寒草地和内蒙古高原温带草地主要双子叶植物叶片解剖特征的比较研究[J].中国科学:生命科学,2012,42(2):158-172.MA J J,JI C J,HAN M,et al.Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands[J].Scientia Sinica(Vitae)2012,42(2):158-172.
[26]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.
[27]ROSSATTO D R,KOLB R M,FRANCO A C.Leaf anatomy is associated with the type of growth form in Neotropical savanna plants[J].Botany,2015,93(8):507-518.
[28]严昌荣,韩兴国,陈灵芝.北京山区落叶阔叶林优势种叶片特点及其生理生态特性[J].生态学报,2000,20(1):53-60.YAN C R,HAN X G,CHEN L Z.The relationship between the ecophysiological feature and leaf characteristics of some woody plants in Beijing mountain zone[J].Acta Ecologica Sinica,2000,20(1):53-60.
[29]郭婧宇,赵杏花,左合君.柠条锦鸡儿小叶解剖结构的环境分异性研究[J].西北植物学报,2017,37(12):2 396-2 402.GUO J Y,ZHAO X H,ZUO H J.Environmental different characteristic study on anatomical structure of Caragana korshinskii leaflet[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(12):2 396-2 402.
[30]BINKS O,MEIR P,et al.Plasticity in leaf-level water relations of tropical rainforest trees in response to experimental drought[J].The New Phytologist,2016,211(2):477-488.
[31]SCOFFONI C,KUNKLE J,PASQUET-KOK J,et al.Light-induced plasticity in leaf hydraulics,venation,anatomy,and gas exchange in ecologically diverse Hawaiian lobeliads[J].The New Phytologist,2015,207(1):43-58.
[2]ACKERLY D,KNIGHT C,WEISS S,et al.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:Contrasting patterns in species level and community level analyses[J].Oecologia,2002,130(3):449-457.
[3]YIN Q L,WANG L,LEI M L,et al.The relationships between leaf economics and hydraulic traits of woody plants depend on water availability[J].The Science of the Total Environment,2018,621:245-252.
[4]BUCKLEY T N,JOHN G P,SCOFFONI C,et al.How does leaf anatomy influence water transport outside the xylem[J].Plant Physiology,2015,168(4):1 616-1 635.
[5]TOMS M,FLEXAS J,et al.Importance of leaf anatomy in determining mesophyll diffusion conductance to CO2 across species:Quantitative limitations and scaling up by models[J].Journal of Experimental Botany,2013,64(8):2 269-2 281.
[6]XIONG D L,FLEXAS J,YU T T,et al.Leaf anatomy mediates coordination of leaf hydraulic conductance and mesophyll conductance to CO2in Oryza[J].The New Phytologist,2017,213(2):572-583.
[7]陈庆诚,孙仰文,张国樑.疏勒河中、下游植物群落优势种生态-形态、解剖特性的初步研究[J].兰州大学学报,1961,(3):65-100.CHEN Q C,SUN Y W,ZHANG G L.A preliminary study on the morphology and anatomy of the dominants of the vegetations distributing along the middle and low drainage basin of the Shor-Ler River based on the view point of ecology[J].Journal of Lanzhou University,1961,(3):65-100.
[8]HE N P,LIU C C,TIAN M,et al.Variation in leaf anatomical traits from tropical to cold-temperate forests and linkage to ecosystem functions[J].Functional Ecology,2018,32(1):10-19.
[9]陈莹婷,许振柱.植物叶经济谱的研究进展[J].植物生态学报,2014,38(10):1 135-1 153.CHEN Y T,XU Z Z.Review on research of leaf economics spectrum[J].Chinese Journal of Plant Ecology,2014,38(10):1 135-1 153.
[10]FU B J,LIU Y,LY H,et al.Assessing the soil erosion control service of ecosystems change in the Loess Plateau of China[J].Ecological Complexity,2011,8(4):284-293.
[11]杜华栋.黄土丘陵沟壑区优势植物对不同侵蚀环境的适应研究---从群落、个体、组织到生理[D].北京:中国科学院大学,2013.
[12]杜华栋,焦菊英,寇萌,等.黄土高原先锋种猪毛蒿叶片形态解剖与生理特征对立地的适应性[J].生态学报,2016,36(10):2 914-2 925.DU H D,JIAO J Y,KOU M,et al.Adaptability of foliar morphological,anatomical,and physiological characteristics of the pioneer species Artemisia scoparia growing in a hillygully Loess Region at different slope sites[J].Acta Ecologica Sinica,2016,36(10):2 914-2 925.
[13]杜华栋,徐翠红,刘萍,等.陕北黄土高原优势植物叶片解剖结构的生态适应性[J].西北植物学报,2010,30(2):293-300.DU H D,XU C H,LIU P,et al.Foliar anatomical structures and ecological adaptabilities of dominant plants in the north Shaanxi Loess Plateau[J].Acta Botanica Boreali-Occidentalia Sinica,2010,30(2):293-300.
[14]王勇,梁宗锁,龚春梅,等.干旱胁迫对黄土高原4种蒿属植物叶形态解剖学特征的影响[J].生态学报,2014,34(16):4 535-4 548.WANG Y,LIANG Z S,GONG C M,et al.Effect of drought on leaf anatomical characteristics of four Artemisia species in the Loess Plateau[J].Acta Ecologica Sinica,2014,34(16):4 535-4 548.
[15]杨超,梁宗锁.陕北撂荒地上优势蒿类叶片解剖结构及其生态适应性[J].生态学报,2008,28(10):4 732-4 738.YANG C,LIANG Z S.Foliar anatomical structures and ecological adaptabilities of dominant artemisia species of early sere of succession on arable old land after being abandoned in Loess Hilly Region[J].Acta Ecologica Sinica,2008,28(10):4 732-4 738.
[16]KOU M,JIAO J Y,YIN Q L,et al.Successional trajectory over 10years of vegetation restoration of abandoned slope croplands in the hill-gully region of the Loess Plateau[J].Land Degradation&Development,2016,27(4):919-932.
[17]中国科学院黄土高原综合科学考察队.黄土高原地区土壤侵蚀区域特征及其治理途径[M].北京:中国科学技术出版社,1991.
[18]王义风.黄土高原地区植被资源及其合理利用[M].北京:科学技术出版社,1991.
[19]李和平.植物显微技术[M].2版.北京:科学出版社,2009.
[20]龚春梅,宁蓬勃,王根轩,等.C3和C4植物光合途径的适应性变化和进化[J].植物生态学报,2009,33(1):206-221.GONG C M,NING P B,WANG G X,et al.A review of adaptable variations and evolution of photosynthetic carbon assimilating pathway in C3and C4plants[J].Chinese Journal of Plant Ecology,2009,33(1):206-221.
[21]李正理.旱生植物的形态和结构[J].生物学通报,1981,16(4):9-12.LI Z L.The structure and morphology of xeromorphic[J].Bulletin of Biology,1981,16(4):9-12.
[22]黄振英,吴鸿,胡正海.30种新疆沙生植物的结构及其对沙漠环境的适应[J].植物生态学报,1997,21(6):521-530.HUANG Z Y,WU H,HU Z H.The structures of 30species of psammophytes and their adaptation to the sandy desert environment in Xinjiang[J].Acta Phytoecologica Sinica,1997,21(6):521-530.
[23]穆兴民.试论黄土区旱地土壤水资源的地带性与非地带性[J].土壤学报,1999,36(2):237-244.MU X M.An exploratory research on the geo-zonal and unzonal distributions of soil water in loess region,China[J].Acta Pedologica Sinica,1999,36(2):237-244.
[24]王力,邵明安,侯庆春.土壤干层量化指标初探[J].水土保持学报,2000,14(4):87-90.WANG L,SHAO M A,HOU Q C.Preliminary research on measured indexes of dried soil layer[J].Journal of Soil and Water Conservation,2000,14(4):87-90.
[25]马建静,吉成均,韩梅,等.青藏高原高寒草地和内蒙古高原温带草地主要双子叶植物叶片解剖特征的比较研究[J].中国科学:生命科学,2012,42(2):158-172.MA J J,JI C J,HAN M,et al.Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands[J].Scientia Sinica(Vitae)2012,42(2):158-172.
[26]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.
[27]ROSSATTO D R,KOLB R M,FRANCO A C.Leaf anatomy is associated with the type of growth form in Neotropical savanna plants[J].Botany,2015,93(8):507-518.
[28]严昌荣,韩兴国,陈灵芝.北京山区落叶阔叶林优势种叶片特点及其生理生态特性[J].生态学报,2000,20(1):53-60.YAN C R,HAN X G,CHEN L Z.The relationship between the ecophysiological feature and leaf characteristics of some woody plants in Beijing mountain zone[J].Acta Ecologica Sinica,2000,20(1):53-60.
[29]郭婧宇,赵杏花,左合君.柠条锦鸡儿小叶解剖结构的环境分异性研究[J].西北植物学报,2017,37(12):2 396-2 402.GUO J Y,ZHAO X H,ZUO H J.Environmental different characteristic study on anatomical structure of Caragana korshinskii leaflet[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(12):2 396-2 402.
[30]BINKS O,MEIR P,et al.Plasticity in leaf-level water relations of tropical rainforest trees in response to experimental drought[J].The New Phytologist,2016,211(2):477-488.
[31]SCOFFONI C,KUNKLE J,PASQUET-KOK J,et al.Light-induced plasticity in leaf hydraulics,venation,anatomy,and gas exchange in ecologically diverse Hawaiian lobeliads[J].The New Phytologist,2015,207(1):43-58.