沙生植物沙鞭叶表皮微形态特征及其生态适应性研究
|
摘要
利用光镜对沙生植物沙鞭50个代表种群的叶表皮微形态特征进行了观测、统计和分析,总结了代表种群叶表皮结构的异同,系统探讨了叶表皮微形态特征的分类学及生态适应性意义。结果表明,沙鞭叶表皮由长细胞、短细胞、气孔器细胞和刺毛组成,无微毛、大毛和乳突,表现为典型的狐茅型;脉间长细胞形态和壁波曲程度、短细胞形态和分布式样、气孔器体积大小与分布及副卫细胞形状、以及刺毛形态等在种群间具有明显差异,可将其鉴分为两大类。同时,沙鞭种群叶表皮气孔密度随环境干旱程度的增加而增加,第一类气孔密度大、第二类气孔密度小,这种结构有利于减少其体内水分的散失,进而更好地适应干旱环境,是对荒漠环境的一种微观形态解剖特征性适应。
We comprehensively observed and analyzed the micromorphological characteristics of leaf epidermises from 50 representative populations of Psammochloa villosa through the light microscopic. We summarized the similarities and differences of leaf epidermis structures,and then systematically discussed the significance of taxonomy and ecological adaptability about micromorphological characteristics of leaf epidermises. The leaf epidermis was composed of long-cells,short-cells,stomatal cells and prickle-hairs,but they had no micro-hair,macro-hair and papillae,which belongs to the typical festucoid type. The obviously different characters,such as the shape of long-cells and curve extent of cell walls,form and distribution pattern of short-cells,volume and distribution of stomas,shape of subsidiary cells and pattern of prickle hair,existed among populations of P. villosa. They can delimitate P. villosa into two groups. Meanwhile,the stomatal density of leaf epidermises in P. villosa increases with the heavier environmental drought. The stomatal density of groupⅠ is bigger that of group Ⅱ. This structure is very beneficial for P. villosa to reduce the loss of water,whichpossibly makes it adapt to arid deserts. Thus,it should be a kind of micromorphologically anatomical adaptation to arid deserts.
We comprehensively observed and analyzed the micromorphological characteristics of leaf epidermises from 50 representative populations of Psammochloa villosa through the light microscopic. We summarized the similarities and differences of leaf epidermis structures,and then systematically discussed the significance of taxonomy and ecological adaptability about micromorphological characteristics of leaf epidermises. The leaf epidermis was composed of long-cells,short-cells,stomatal cells and prickle-hairs,but they had no micro-hair,macro-hair and papillae,which belongs to the typical festucoid type. The obviously different characters,such as the shape of long-cells and curve extent of cell walls,form and distribution pattern of short-cells,volume and distribution of stomas,shape of subsidiary cells and pattern of prickle hair,existed among populations of P. villosa. They can delimitate P. villosa into two groups. Meanwhile,the stomatal density of leaf epidermises in P. villosa increases with the heavier environmental drought. The stomatal density of groupⅠ is bigger that of group Ⅱ. This structure is very beneficial for P. villosa to reduce the loss of water,whichpossibly makes it adapt to arid deserts. Thus,it should be a kind of micromorphologically anatomical adaptation to arid deserts.
引文
1.蔡联炳,王世金,李建华.中国主要禾本科植物叶片表皮细胞、表皮附属物的常见类型及其分类价值的初步探讨[C].高原生物学集刊,1991(10):1-14.Cai L B,Wang S J,LI J H.Primary study on the classification value and the ordinary type of the attachment for the leaf cell epidermal is of major Poaceae in China[C].The Journal Polymer Published in Plateau Biology,1991,(10):1-14.
2 .Prat H.La systématique des Graminées[J].Ann Sci Nat Bot,1936,10(18):165-258.
3 .Metcalfe C R.Anatomy of the monocotyledons:Volume 1Gramineae[M].London:Oxford Clarendon Press,1960:1
4 .Clifford H T,Watson L.Identifying grasses:data,methods and illustrations[M].Brisbane:University of Queensland Press,1977:1-141.
5 .陈守良,金岳杏,吴竹君,等.叶表皮细胞结构在国产狗尾草属(Setaria Beauv.)分组水平上的应用[J].植物研究,1985,5(2):105-112.Chen S L,Jin Y X,Wu Z J,et al.The study leaf epidermis at the sectional level on Chinese Setaria[J].Bulletin of Botanical Research,1985,5(2):105-112.
6 .蔡联炳,郭延平.禾本科植物叶片表皮结构细胞主要类型的演化与系统分类和发育途径的探讨[J].西北植物学报,1995,15(4):323-335.Cai L B,Guo Y P.Studies on constituent cells of leaf epidermis,systematics and phylogenetic path of the family Poaceae[J].Acta Botanica Boreali-Occidentalia Sinica,1995,15(4):323-335.
7 .蔡联炳,郭延平.禾本科植物叶片表皮结构细胞主要类型的演化与系统分类和发育途径的探讨(续)[J].西北植物学报,1996,16(1):65-72.Cai L B,Guo Y P.Studies on constituent cells of leaf epidermis,systematics and phylogenetic path of the family Poaceae(Continued)[J].Acta Botanica Boreali-Occidentalia Sinica,1996,16(1):65-72.
8 .蔡联炳.禾本科叶片表皮结构细胞的组合式样及其分类学意义[J].植物研究,1999,19(4):415-427.Cai L B.Combinative patterns of constituent cells of leaf epidermis of Poaceae and their taxonomic significance[J].Bulletin of Botanical Research,1999,19(4):415-427.
9 .苏旭,刘玉萍,李兆孟,等.基于形态和叶表皮微形态特征试论西藏固沙草的归并[J].植物研究,2015,35(3):321-326.Su X,Liu Y P,Li Z M,et al.Reduction on Orinus tibeticus based on the external morphology and micromorphological characteristics of leaf epidermis[J].Bulletin of Botanical Research,2015,35(3):321-326.
10 .马海英,彭华,王跃华.禾本科广义拂子茅属的叶表皮形态研究[J].植物分类学报,2006,44(4):371-392.Ma H Y,Peng H,Wang Y H.Morphology of leaf epidermis of Calamagrostis s.l.(Poaceae:Pooideae)in China[J].Acta Phytotaxonomica Sinica,2006,44(4):371-392.
11 .李兆孟,周勇辉,刘玉萍,等.基于形态和叶表皮微形态特征试论国产赖草属属下类群间的系统关系[J].植物研究,2016,36(1):10-25.Li Z M,Zhou Y H,Liu Y P,et al.Biosystematic relationships among taxa below Leymus(Poaceae)from China based on the external morphology and micromorphological characteristics of leaf epidermis[J].Bulletin of Botanical Research,2016,36(1):10-25.
12 .刘瑛心.中国沙漠植物志:第1卷[M].北京:科学出版社,1985:34.Liu Y X.Flora in desertis reipublicae populorum sinarum(Tomus 1)[M].Beijing:Science Press,1985:34.
13 .郭本兆.中国植物志:第9卷[M].北京:科学出版社,1987:309.Kuo P C.Flora Reipublicae Popularis Sinicae(Tomus 9)[M].Beijing:Science Press,1987:309.
14 .马毓泉.内蒙古植物志:第5卷[M].2版.呼和浩特:内蒙古人民出版社,1994:115-152.Ma Y Q.Flora Inner Mongolica(Tomus 5)[M].2nd ed.Hohhot:Inner Mongolia People’s Publishing House,1994:115-152.
15 .Wu Z Y,Raven P H,Hong D Y,et al.Flora of China(Vol.22)[M].Beijing:Science Press and St.Louis:Missouri Botanical Garden Press,2006.
16 .陈文俐.中国野青茅属的分类学修订[D].北京:中国科学院植物研究所,2001.Chen W L.Revision of the genus Deyeuxia(Poaceae)in China[D].Beijing:The Institute of Botany,Chinese Academy of Sciences,2001.
17 .董鸣,阿拉腾宝,邢雪荣,等.根茎禾草沙鞭的克隆基株及分株种群特征[J].植物生态学报,1999,23(4):302-310.Dong M,Alateng B,Xing X R,et al.Genet features and ramet population features in the rhizomatous grass species Psammochloa villosa[J].Acta Phytoecologica Sinica,1999,23(4):302-310.
18 .董鸣.切断根茎对根茎禾草沙鞭和赖草克隆生长的影响[J].植物学报,1999,41(2):194-198.Dong M.Effects of severing rhizome on clonal growth in rhizomatous grass species Psammochloa villosa and Leymus secalinus[J].Acta Botanica Sinica,1999,41(2):194-198.
19 .王可青,葛颂,董鸣.根茎禾草沙鞭的等位酶变异及克隆多样性[J].植物学报,1999,41(5):537-540.Wang K Q,Ge S,Dong M.Allozyme variance and clonal diversity in the rhizomatous grass Psammochloa villosa(Gramineae)[J].Acta Botanica Sinica,1999,41(5):537-540.
20 .陈玉福,于飞海,张称意,等.根茎禾草沙鞭的克隆生长在毛乌素沙地斑块动态中的作用[J].生态学报,2001,21(11):1745-1750.Chen Y F,Yu F H,Zhang C Y,et al.Role of clonal growth of the rehizomatous grass Psammochloa villosa in patch dynamics of Mu Us sandy land[J].Acta Ecologica Sinica,2001,21(11):1745-1750.
21 .黄振英.鄂尔多斯高原固沙禾草沙鞭种子休眠和萌发与环境的关系[J].西北植物学报,2003,23(7):1128-1133.Huang Z Y.Adaptation strategies of seed dormancy and germination of Psammochloa villosa,a sand dune grass inhabiting Ordos Plateau,China[J].Acta Botanica BorealiOccidentalia Sinica,2003,23(7):1128-1133.
22 .黄振英,董鸣,张淑敏.沙鞭(禾本科)种子在沙丘上的萌发策略及幼苗的耐干燥特性[J].生态学报,2005,25(2):298-303.Huang Z Y,Dong M,Zhang S M.Strategies of seed germination on sand dune and seedling desiccation tolerance,of Psammochloa villosa(Poaceae)[J].Acta Ecologica Sinica,2005,25(2):298-303.
23 .朱雅娟,董鸣,黄振英.沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响[J].植物生态学报,2005,29(5):730-739.Zhu Y J,Dong M,Huang Z Y.Effects of sand burial and seed size on seed germination and seedling emergence of Psammochloa villosa[J].Acta Phytoecologica Sinica,2005,29(5):730-739.
24 .徐翥骍,贺学礼,郭辉娟,等.内蒙古农牧交错区沙鞭和羊柴AM真菌侵染及其土壤因子[J].河北农业大学学报,2011,34(1):56-61.Xu Z X,He X L,Guo H J,et al.AM fungi colonization and soil factors of Psammochloa villosa(Trin.)Bor and Hedysalum laeve Min.in farming-pastoral area of Inner Mongolia[J].Journal of Agricultural University of Hebei,2011,34(1):56-61.
25 .马成仓,高玉葆,王金龙,等.内蒙古高原甘蒙锦鸡儿光合作用和水分代谢的生态适应性研究[J].植物生态学报,2004,28(3):305-311.Ma C C,Gao Y B,Wang J L,et al.Ecological adaptation of Caragana opulens on the inner Mongolia Plateau:photosynthesis and water metabolism[J].Acta Phytoecologica Sinica,2004,28(3):305-311.
26 .周云龙.植物生物学[M].2版.北京:高等教育出版社,2004.Zhou Y L.Plant biology[M].2nd ed.Beijing:Higher Education Press,2004.
27 .李长复.叶子表面的适应性能[J].生物学通报,1985(9):2+5-7.Li C F.Adaptability of leaf surface[J].Bulletin of Biology,1985(9):2+5-7.
28 .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,2005,22(S1):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,2005,22(S1):118-127.
29 .刘玉冰,李新荣,李蒙蒙,等.中国干旱半干旱区荒漠植物叶片(或同化枝)表皮微形态特征[J].植物生态学报,2016,40(11):1189-1207.Liu Y B,Li X R,Li M M,et al.Leaf(or assimilation branch)epidermal micromorphology of desert plant in arid and semiarid areas of China[J].Chinese Journal of Plant Ecology,2016,40(11):1189-1207.
30 .Barthlott W.Epidermal and seed surface characters of plants:systematic applicability and some evolutionary aspects[J].Nordic Journal of Botany,1981,1(3):345-355.
31 .Casson S A,Hetherington A M.Environmental regulation of stomatal development[J].Current Opinion in Plant Biology,2010,13(1):90-95.
32 .Ghahremaninejad F,Khalili Z,Maassoumi A A,et al.Leaf epidermal features of Salix species(Salicaceae)and their systematic significance[J].American Journal of Botany,2012,99(4):769-777.
33 .王勋陵,马骥.从旱生植物叶结构探讨其生态适应的多样性[J].生态学报,1999,19(6):787-792.Wang X L,Ma J.A study on leaf structure and the diversity of xerophytes ecology adaptation[J].Acta Ecologica Sinica,1999,19(6):787-792.
2 .Prat H.La systématique des Graminées[J].Ann Sci Nat Bot,1936,10(18):165-258.
3 .Metcalfe C R.Anatomy of the monocotyledons:Volume 1Gramineae[M].London:Oxford Clarendon Press,1960:1
4 .Clifford H T,Watson L.Identifying grasses:data,methods and illustrations[M].Brisbane:University of Queensland Press,1977:1-141.
5 .陈守良,金岳杏,吴竹君,等.叶表皮细胞结构在国产狗尾草属(Setaria Beauv.)分组水平上的应用[J].植物研究,1985,5(2):105-112.Chen S L,Jin Y X,Wu Z J,et al.The study leaf epidermis at the sectional level on Chinese Setaria[J].Bulletin of Botanical Research,1985,5(2):105-112.
6 .蔡联炳,郭延平.禾本科植物叶片表皮结构细胞主要类型的演化与系统分类和发育途径的探讨[J].西北植物学报,1995,15(4):323-335.Cai L B,Guo Y P.Studies on constituent cells of leaf epidermis,systematics and phylogenetic path of the family Poaceae[J].Acta Botanica Boreali-Occidentalia Sinica,1995,15(4):323-335.
7 .蔡联炳,郭延平.禾本科植物叶片表皮结构细胞主要类型的演化与系统分类和发育途径的探讨(续)[J].西北植物学报,1996,16(1):65-72.Cai L B,Guo Y P.Studies on constituent cells of leaf epidermis,systematics and phylogenetic path of the family Poaceae(Continued)[J].Acta Botanica Boreali-Occidentalia Sinica,1996,16(1):65-72.
8 .蔡联炳.禾本科叶片表皮结构细胞的组合式样及其分类学意义[J].植物研究,1999,19(4):415-427.Cai L B.Combinative patterns of constituent cells of leaf epidermis of Poaceae and their taxonomic significance[J].Bulletin of Botanical Research,1999,19(4):415-427.
9 .苏旭,刘玉萍,李兆孟,等.基于形态和叶表皮微形态特征试论西藏固沙草的归并[J].植物研究,2015,35(3):321-326.Su X,Liu Y P,Li Z M,et al.Reduction on Orinus tibeticus based on the external morphology and micromorphological characteristics of leaf epidermis[J].Bulletin of Botanical Research,2015,35(3):321-326.
10 .马海英,彭华,王跃华.禾本科广义拂子茅属的叶表皮形态研究[J].植物分类学报,2006,44(4):371-392.Ma H Y,Peng H,Wang Y H.Morphology of leaf epidermis of Calamagrostis s.l.(Poaceae:Pooideae)in China[J].Acta Phytotaxonomica Sinica,2006,44(4):371-392.
11 .李兆孟,周勇辉,刘玉萍,等.基于形态和叶表皮微形态特征试论国产赖草属属下类群间的系统关系[J].植物研究,2016,36(1):10-25.Li Z M,Zhou Y H,Liu Y P,et al.Biosystematic relationships among taxa below Leymus(Poaceae)from China based on the external morphology and micromorphological characteristics of leaf epidermis[J].Bulletin of Botanical Research,2016,36(1):10-25.
12 .刘瑛心.中国沙漠植物志:第1卷[M].北京:科学出版社,1985:34.Liu Y X.Flora in desertis reipublicae populorum sinarum(Tomus 1)[M].Beijing:Science Press,1985:34.
13 .郭本兆.中国植物志:第9卷[M].北京:科学出版社,1987:309.Kuo P C.Flora Reipublicae Popularis Sinicae(Tomus 9)[M].Beijing:Science Press,1987:309.
14 .马毓泉.内蒙古植物志:第5卷[M].2版.呼和浩特:内蒙古人民出版社,1994:115-152.Ma Y Q.Flora Inner Mongolica(Tomus 5)[M].2nd ed.Hohhot:Inner Mongolia People’s Publishing House,1994:115-152.
15 .Wu Z Y,Raven P H,Hong D Y,et al.Flora of China(Vol.22)[M].Beijing:Science Press and St.Louis:Missouri Botanical Garden Press,2006.
16 .陈文俐.中国野青茅属的分类学修订[D].北京:中国科学院植物研究所,2001.Chen W L.Revision of the genus Deyeuxia(Poaceae)in China[D].Beijing:The Institute of Botany,Chinese Academy of Sciences,2001.
17 .董鸣,阿拉腾宝,邢雪荣,等.根茎禾草沙鞭的克隆基株及分株种群特征[J].植物生态学报,1999,23(4):302-310.Dong M,Alateng B,Xing X R,et al.Genet features and ramet population features in the rhizomatous grass species Psammochloa villosa[J].Acta Phytoecologica Sinica,1999,23(4):302-310.
18 .董鸣.切断根茎对根茎禾草沙鞭和赖草克隆生长的影响[J].植物学报,1999,41(2):194-198.Dong M.Effects of severing rhizome on clonal growth in rhizomatous grass species Psammochloa villosa and Leymus secalinus[J].Acta Botanica Sinica,1999,41(2):194-198.
19 .王可青,葛颂,董鸣.根茎禾草沙鞭的等位酶变异及克隆多样性[J].植物学报,1999,41(5):537-540.Wang K Q,Ge S,Dong M.Allozyme variance and clonal diversity in the rhizomatous grass Psammochloa villosa(Gramineae)[J].Acta Botanica Sinica,1999,41(5):537-540.
20 .陈玉福,于飞海,张称意,等.根茎禾草沙鞭的克隆生长在毛乌素沙地斑块动态中的作用[J].生态学报,2001,21(11):1745-1750.Chen Y F,Yu F H,Zhang C Y,et al.Role of clonal growth of the rehizomatous grass Psammochloa villosa in patch dynamics of Mu Us sandy land[J].Acta Ecologica Sinica,2001,21(11):1745-1750.
21 .黄振英.鄂尔多斯高原固沙禾草沙鞭种子休眠和萌发与环境的关系[J].西北植物学报,2003,23(7):1128-1133.Huang Z Y.Adaptation strategies of seed dormancy and germination of Psammochloa villosa,a sand dune grass inhabiting Ordos Plateau,China[J].Acta Botanica BorealiOccidentalia Sinica,2003,23(7):1128-1133.
22 .黄振英,董鸣,张淑敏.沙鞭(禾本科)种子在沙丘上的萌发策略及幼苗的耐干燥特性[J].生态学报,2005,25(2):298-303.Huang Z Y,Dong M,Zhang S M.Strategies of seed germination on sand dune and seedling desiccation tolerance,of Psammochloa villosa(Poaceae)[J].Acta Ecologica Sinica,2005,25(2):298-303.
23 .朱雅娟,董鸣,黄振英.沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响[J].植物生态学报,2005,29(5):730-739.Zhu Y J,Dong M,Huang Z Y.Effects of sand burial and seed size on seed germination and seedling emergence of Psammochloa villosa[J].Acta Phytoecologica Sinica,2005,29(5):730-739.
24 .徐翥骍,贺学礼,郭辉娟,等.内蒙古农牧交错区沙鞭和羊柴AM真菌侵染及其土壤因子[J].河北农业大学学报,2011,34(1):56-61.Xu Z X,He X L,Guo H J,et al.AM fungi colonization and soil factors of Psammochloa villosa(Trin.)Bor and Hedysalum laeve Min.in farming-pastoral area of Inner Mongolia[J].Journal of Agricultural University of Hebei,2011,34(1):56-61.
25 .马成仓,高玉葆,王金龙,等.内蒙古高原甘蒙锦鸡儿光合作用和水分代谢的生态适应性研究[J].植物生态学报,2004,28(3):305-311.Ma C C,Gao Y B,Wang J L,et al.Ecological adaptation of Caragana opulens on the inner Mongolia Plateau:photosynthesis and water metabolism[J].Acta Phytoecologica Sinica,2004,28(3):305-311.
26 .周云龙.植物生物学[M].2版.北京:高等教育出版社,2004.Zhou Y L.Plant biology[M].2nd ed.Beijing:Higher Education Press,2004.
27 .李长复.叶子表面的适应性能[J].生物学通报,1985(9):2+5-7.Li C F.Adaptability of leaf surface[J].Bulletin of Biology,1985(9):2+5-7.
28 .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,2005,22(S1):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,2005,22(S1):118-127.
29 .刘玉冰,李新荣,李蒙蒙,等.中国干旱半干旱区荒漠植物叶片(或同化枝)表皮微形态特征[J].植物生态学报,2016,40(11):1189-1207.Liu Y B,Li X R,Li M M,et al.Leaf(or assimilation branch)epidermal micromorphology of desert plant in arid and semiarid areas of China[J].Chinese Journal of Plant Ecology,2016,40(11):1189-1207.
30 .Barthlott W.Epidermal and seed surface characters of plants:systematic applicability and some evolutionary aspects[J].Nordic Journal of Botany,1981,1(3):345-355.
31 .Casson S A,Hetherington A M.Environmental regulation of stomatal development[J].Current Opinion in Plant Biology,2010,13(1):90-95.
32 .Ghahremaninejad F,Khalili Z,Maassoumi A A,et al.Leaf epidermal features of Salix species(Salicaceae)and their systematic significance[J].American Journal of Botany,2012,99(4):769-777.
33 .王勋陵,马骥.从旱生植物叶结构探讨其生态适应的多样性[J].生态学报,1999,19(6):787-792.Wang X L,Ma J.A study on leaf structure and the diversity of xerophytes ecology adaptation[J].Acta Ecologica Sinica,1999,19(6):787-792.