胡杨异形叶性状与其个体发育的关系
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摘要
【目的】叶片是植物适应环境的重要器官,叶片结构性状的改变影响植物光合作用等生理过程,探讨胡杨叶形变化规律及其影响因素对理解其适应特性具有重要意义。【方法】为明确胡杨异形叶结构性状特征与其个体发育之间的关系,在额济纳人工胡杨林内调查不同树龄、繁殖期与非繁殖期胡杨树冠下层叶片的长与宽以分析其叶形的变化规律。【结果】胡杨异形叶形成是一个渐变的过程,随着树龄增加,胡杨叶长减小,叶宽增加,叶长宽比减小;树龄对叶片长、宽和长宽比变异的解释率分别为53. 1%、38. 6%和66. 1%。繁殖期对胡杨叶形变化具有显著影响,6年生胡杨繁殖期叶长和叶长宽比显著小于非繁殖期叶片,而叶宽显著大于非繁殖期叶片。非繁殖期的胡杨叶形则随树龄增加由条形叶向卵形叶逐渐转变。胸径与胡杨叶片形态特征显著相关,随着胸径增大,胡杨叶长与叶长宽比减小,叶宽增大;在一元回归分析中,胸径对胡杨叶片形态特征的解释率均大于0. 5(R2> 0. 5,P <0. 000 1),而在多元回归中,胸径单独的解释率为8. 6%(叶长)、13. 1%(叶宽)、3. 4%(叶长宽比);此外,胸径与树龄对胡杨叶片结构性状特征具有显著的交互影响。【结论】胡杨异形叶的发生与其个体生长发育之间具有密切关系,随着树木的生长,胡杨叶形由条形叶向卵形、阔卵形叶转变以适应干旱高温的环境;此外,繁殖期加速胡杨叶形的转变,可能是胡杨应对繁殖期能量需求的适应性策略。
[Objective] Leaf is the most adaptable plant organ responding to the environment. Leaf structure influences plant physiological processes,such as photosynthesis. Thus,it is extremely important to understand the changes in leaf structure and underly factors for exploring the strategy of plant adaptation. [Method] To detect the relationship between the leaf structural traits of Populus euphratica and its individual development stage,we investigated the leaf length and width of P. euphratica to calculate leaf shape changes during reproduction and non-reproduction period at different ages in Ejina artificial P. euphratica forest of Inner Mongolia. [Result]The results showed that the leaf length and the ratio of leaf length to leaf width decreased with increasing tree age,while the leaf width increased. Treeage accounted for 53. 1%,38. 6% and 66. 1% of variations in the leaf length,leaf width and the ratio of leaf length to width,respectively. The leaf shape changes were significantly influenced by reproduction period. Leaf length and ratio of the length to width in reproduction period were significantly smaller while leaf width was larger than those in non-reproduction period. However,the leaves of P. euphratica during non-reproduction period gradually changed from lanceolate to broad-ovate with increased tree age. The DBH was significantly correlated with the leaf traits,which was negatively correlated with the leaf length and the ratio of leaf length to leaf width versus positively correlated with the leaf width. The DBH accounted for most of the variations in the leaf traits by one-dimensional regression analysis( R2> 0. 5,P < 0. 000 1). But DBH accounted for a lower portion of the overall variations of leaf traits based on multiple linear models( leaf length 8. 6%,leaf width 13. 1%,the ratio of leaf length to width 3. 4%).Furthermore, we detected significant interactive effects between DBH and age on the leaf traits.[Conclusion]In conclusion,the leaf traits of P. euphratica were significantly correlated with the tree growth and development. The leaf shape changed from lanceolate to broad-ovate with tree growth to adapt the desert environment. We infer P. euphratica accelerates the leaf shape changes to provide sufficient energy for reproduction which may be an adaptive strategy of P. euphratica to environmental conditions.
[Objective] Leaf is the most adaptable plant organ responding to the environment. Leaf structure influences plant physiological processes,such as photosynthesis. Thus,it is extremely important to understand the changes in leaf structure and underly factors for exploring the strategy of plant adaptation. [Method] To detect the relationship between the leaf structural traits of Populus euphratica and its individual development stage,we investigated the leaf length and width of P. euphratica to calculate leaf shape changes during reproduction and non-reproduction period at different ages in Ejina artificial P. euphratica forest of Inner Mongolia. [Result]The results showed that the leaf length and the ratio of leaf length to leaf width decreased with increasing tree age,while the leaf width increased. Treeage accounted for 53. 1%,38. 6% and 66. 1% of variations in the leaf length,leaf width and the ratio of leaf length to width,respectively. The leaf shape changes were significantly influenced by reproduction period. Leaf length and ratio of the length to width in reproduction period were significantly smaller while leaf width was larger than those in non-reproduction period. However,the leaves of P. euphratica during non-reproduction period gradually changed from lanceolate to broad-ovate with increased tree age. The DBH was significantly correlated with the leaf traits,which was negatively correlated with the leaf length and the ratio of leaf length to leaf width versus positively correlated with the leaf width. The DBH accounted for most of the variations in the leaf traits by one-dimensional regression analysis( R2> 0. 5,P < 0. 000 1). But DBH accounted for a lower portion of the overall variations of leaf traits based on multiple linear models( leaf length 8. 6%,leaf width 13. 1%,the ratio of leaf length to width 3. 4%).Furthermore, we detected significant interactive effects between DBH and age on the leaf traits.[Conclusion]In conclusion,the leaf traits of P. euphratica were significantly correlated with the tree growth and development. The leaf shape changed from lanceolate to broad-ovate with tree growth to adapt the desert environment. We infer P. euphratica accelerates the leaf shape changes to provide sufficient energy for reproduction which may be an adaptive strategy of P. euphratica to environmental conditions.
引文
[1]刘金环,曾德慧,Don Koo Lee.科尔沁沙地东南部地区主要植物叶片性状及其相互关系[J].生态学杂志,2006,25(8):921-925.Liu J H, Zeng D H, Lee D K. Leaf traits and their interrelationships of main plant species in southeast Horqin Sandy Land[J]. Chinese Journal of Ecology,2006,25(8):921--925.
[2] Kikuzawa K. Leaf phenology as an optimal strategy for carbon gain in plants[J]. Canadian Journal of Botany,1995,73(2):158-163.
[3]刘晓娟,马克平.植物功能性状研究进展[J].中国科学:生命科学,2015,45(4):325--339.Liu X J,Ma K P. Plant functional traits:concepts,applications and future directions[J]. Scientia China(Life Sciences),2015,45(4):325--339.
[4] Sun L,Zhang B,Wang B,et al. Leaf elemental stoichiometry of Tamarix Lour species in relation to geographic,climatic,soil,and genetic components in China[J]. Ecological Engineering,2017,106:448--457.
[5] Garnier E,Shipley B,Roumet C,et al. A standardized protocol for the determination of specific leaf area and leaf dry matter content[J]. Functional Ecology,2001,15(5):688-695.
[6]赵红洋,李玉霖,王新源,等.科尔沁沙地52种植物叶片性状变异特征研究[J].中国沙漠,2010,30(6):1292--1298.Zhao H Y,Li Y L,Wang X Y,et al. Variations in leaf traits of52 plants in Horqin Sand Land[J]. Journal of Desert Research,2010,30(6):1292-1298.
[7]李善家,苏培玺,张海娜,等.荒漠植物叶片水分和功能性状特征及其相互关系[J].植物生理学报,2013,49(2):153-160.Li S J, Su P X, Zhang H N, et al. Characteristics and relationships of foliar water and leaf functional traits of desert plants[J]. Plant Physiology Journal,2013,49(2):153-160.
[8]王世绩.全球胡杨林的现状及保护和恢复对策[J].世界林业研究,1996(6):38--45.Wang S J. The status, conservation and recovery of global resources of Populus euphradica[J]. World Forestry Research,1996(6):38-45.
[9]周洪华,李卫红,李玉朋,等.基于树木年轮技术的塔里木河下游河岸胡杨林生态需水量研究[J].生态学报,2017,37(22):7576--7584.Zhou H H,Li W H,Li Y P,et al. Estimation of ecological water demand of a desert riparian forest using tree rings of Populus euphratica in the lower Tarim River[J]. Acta Ecologica Sinica,2017,37(22):7576-7584.
[10]陈黎,周玲玲,庄丽,等.塔里木河下游胡杨三种异形叶的光合特性研究[J].北方园艺,2014(18):88--93.Chen L,Zhou L L,Zhuang L,et al. Study on photosynthetic characteristics of the three heteromorphic leaves of Populus euphratica in the lower Tarim River[J]. Northern Horticulture,2014(18):88-93.
[11]李景文.大漠胡杨[J].大自然,2014(6):38--41.Li J W. Populus euphratica in desert[J]. China Nature,2014(6):38-41.
[12]冯梅.胡杨叶形变化与个体径级间的关系研究[C]∥中国植物学会第十五届会员代表大会暨八十周年学术年会论文集.南昌:中国植物学会,2013.Feng M. Relationship between leaf shape change and individual diameter of Populus euphratica[C]∥Proceedings of the 15th member congress and the 80th annual meeting of the Chinese plant society. Nanchang:Chinese plant society,2013.
[13]肖磊,陈宁美,陈悦,等.内蒙古与北京地区胡杨异形叶表皮蜡质及气孔形态显微结构差异[J].中央民族大学学报(自然科学版),2016,25(3):85-91.Xiao L,Chen N M,Chen Y,et al. The difference of cuticle wax crystallization and stoma morphology of lanceolate and broad-ovate leaves of Populus euphratica olive between Ejina Area in Inner Mongolia and Beijing Area[J]. Journal of MUC(Natural Sciences Edition),2016,25(3):85-91.
[14]杨树德,郑文菊,陈国仓,等.胡杨披针形叶与宽卵形叶的超微结构与光合特性的差异[J].西北植物学报,2005,25(1):14-21.Yang S D, Zheng W J, Chen G C, et al. Difference of ultrastructure and photosynthetic characteristics between lanceolate and broad-ovate leaves in Populus euphratica[J]. Acta Botanica Boreali-Occidentalia Sinica,2005,25(1):14-21.
[15]王海珍,韩路,徐雅丽,等.胡杨异形叶光合作用对光强与CO2浓度的响应[J].植物生态学报,2014,38(10):1099-1109.Wang H Z,Han L,Xu Y L,et al. Photosynthetic responses of the heteromorphic leaves in Populus euphratica to light intensity and CO2concentration[J]. Chinese Journal of Plant Ecology,2014,38(10):1099--1109.
[16]李雁玲,张肖,冯梅,等.胡杨(Populus euphratica)异形叶叶片内源激素特征研究[J].塔里木大学学报,2017,29(3):7-13.Li Y L,Zhang X,Fen M,et al. Characteristics of endhormones in leaf blade of Populus euphratica heteromorphic leaves[J]. Journal of Tarim University,2017,29(3):7--13.
[17]王玉丽,焦培培,顾亚亚,等.胡杨叶形变化与POD同工酶的关系研究[J].干旱区资源与环境,2013,27(4):181-186.Wang Y L,Jiao P P,Gu Y Y,et al. Study on the relationship between leaf shape change and POD enzyme of Populus euphratica[J]. Journal of Arid Land Resources and Environment,2013,27(4):181--186.
[18]秦少伟,李才林,郭忠忠,等.参与胡杨异形叶发生的发育相关基因表达模式及作用[J].基因组学与应用生物学,2017,36(11):4785--4792.Qin S W,Li C L,Guo Z Z,et al. Expression patterns and functions of genes related to the development of Populus euphratica Oliv. heteromorphic leaves morphogenesis[J]. Genomics and Applied Biology,2017,36(11):4785--4792.
[19]秦少伟,赵利峰,张娜,等. rot3、rot4、an和spk1在胡杨异形叶发生中的表达模式和作用[J].分子植物育种,2016,14(10):2630--2636.Qin S W,Zhao L F,Zhang N,et al. Expression patterns and effects of rot3, rot4, an and spk1 on the genesis of Populus euphratica Oliv. heteromorphic leaves[J]. Molecular Plant Breeding,2016,14(10):2630--2636.
[20]黄文娟,李志军,杨赵平,等.胡杨异形叶结构型性状及其与胸径关系[J].生态学杂志,2010,29(12):2347-2352.Huang W J, Li Z J, Yang Z P, et al. Heteromorphic leaf structural characteristics and their correlations with diameter at breast height of Populus euphratica[J]. Chinese Journal of Ecology,2010,29(12):2347-2352.
[21]岳宁,郑彩霞.胡杨异形叶蛋白质表达的电泳分析[J].安徽农业科学,2010,38(32):18058-18061.Yue N,Zheng C X. Electrophoresis analysis on the expressed proteinsin heteromorphic leaves of Populus euphratica Oliv.[J].Journal of Anhui Agri Sci,2010,38(32):18058-18061.
[22]苏培玺,张立新,杜明武,等.胡杨不同叶形光合特性、水分利用效率及其对加富CO2的响应[J].植物生态学报,2003,27(1):34--40.Su P X,Zhang L X,Du M W,et al. Photosynthetic character and water use efficiency of different leaf shapes of Populus euphratica and their response to CO2enrichment[J]. Acta Phytoecologica Sinica,2003,27(1):34--40.
[23]黄文娟,李志军,杨赵平,等.胡杨异形叶结构型性状及其相互关系[J].生态学报,2010,30(17):4636-4642.Huang W J,Li Z J,Yang Z P,et al. The structural traits of Populus euphratica heteromorphic leaves and their correlations[J]. Acta Ecologica Sinica,2010,30(17):4636--4642.
[24]刘帅飞,焦培培,李志军.灰叶胡杨异形叶的类型及其时空特征[J].干旱区研究,2016,33(5):1098--1103.Liu S F,Jiao P P,Li Z J. Diversifolious types and spatiotemporal characteristics of Populus pruinosa schrenk[J]. Arid Zone Research,2016,33(5):1098--1103.
[25] Weiner J. Allocation,plasticity and allometry in plants[J].Perspectives in Plant Ecology,Evolution and Systematics,2004,6(4):207--215.
[26] Zheng C, Qiu J, Jiang C, et al. Comparison of stomatal characteristics and photosynthesis of polymorphic Populus euphratica leaves[J]. Frontiers of Forestry in China,2007,2(1):87--93.
[27]赵鹏宇,冯梅,焦培培,等.胡杨不同发育阶段叶片形态解剖学特征及其与胸径的关系[J].干旱区研究,2016,33(5):1071-1080.Zhao P Y, Feng M, Jiao P P, et al. Relationship between morphological or anatomical features of leaves and trunk diameter at breast height at different growing stages of Populus euphratica[J]. Arid Zone Research,2016,33(5):1071--1080.
[2] Kikuzawa K. Leaf phenology as an optimal strategy for carbon gain in plants[J]. Canadian Journal of Botany,1995,73(2):158-163.
[3]刘晓娟,马克平.植物功能性状研究进展[J].中国科学:生命科学,2015,45(4):325--339.Liu X J,Ma K P. Plant functional traits:concepts,applications and future directions[J]. Scientia China(Life Sciences),2015,45(4):325--339.
[4] Sun L,Zhang B,Wang B,et al. Leaf elemental stoichiometry of Tamarix Lour species in relation to geographic,climatic,soil,and genetic components in China[J]. Ecological Engineering,2017,106:448--457.
[5] Garnier E,Shipley B,Roumet C,et al. A standardized protocol for the determination of specific leaf area and leaf dry matter content[J]. Functional Ecology,2001,15(5):688-695.
[6]赵红洋,李玉霖,王新源,等.科尔沁沙地52种植物叶片性状变异特征研究[J].中国沙漠,2010,30(6):1292--1298.Zhao H Y,Li Y L,Wang X Y,et al. Variations in leaf traits of52 plants in Horqin Sand Land[J]. Journal of Desert Research,2010,30(6):1292-1298.
[7]李善家,苏培玺,张海娜,等.荒漠植物叶片水分和功能性状特征及其相互关系[J].植物生理学报,2013,49(2):153-160.Li S J, Su P X, Zhang H N, et al. Characteristics and relationships of foliar water and leaf functional traits of desert plants[J]. Plant Physiology Journal,2013,49(2):153-160.
[8]王世绩.全球胡杨林的现状及保护和恢复对策[J].世界林业研究,1996(6):38--45.Wang S J. The status, conservation and recovery of global resources of Populus euphradica[J]. World Forestry Research,1996(6):38-45.
[9]周洪华,李卫红,李玉朋,等.基于树木年轮技术的塔里木河下游河岸胡杨林生态需水量研究[J].生态学报,2017,37(22):7576--7584.Zhou H H,Li W H,Li Y P,et al. Estimation of ecological water demand of a desert riparian forest using tree rings of Populus euphratica in the lower Tarim River[J]. Acta Ecologica Sinica,2017,37(22):7576-7584.
[10]陈黎,周玲玲,庄丽,等.塔里木河下游胡杨三种异形叶的光合特性研究[J].北方园艺,2014(18):88--93.Chen L,Zhou L L,Zhuang L,et al. Study on photosynthetic characteristics of the three heteromorphic leaves of Populus euphratica in the lower Tarim River[J]. Northern Horticulture,2014(18):88-93.
[11]李景文.大漠胡杨[J].大自然,2014(6):38--41.Li J W. Populus euphratica in desert[J]. China Nature,2014(6):38-41.
[12]冯梅.胡杨叶形变化与个体径级间的关系研究[C]∥中国植物学会第十五届会员代表大会暨八十周年学术年会论文集.南昌:中国植物学会,2013.Feng M. Relationship between leaf shape change and individual diameter of Populus euphratica[C]∥Proceedings of the 15th member congress and the 80th annual meeting of the Chinese plant society. Nanchang:Chinese plant society,2013.
[13]肖磊,陈宁美,陈悦,等.内蒙古与北京地区胡杨异形叶表皮蜡质及气孔形态显微结构差异[J].中央民族大学学报(自然科学版),2016,25(3):85-91.Xiao L,Chen N M,Chen Y,et al. The difference of cuticle wax crystallization and stoma morphology of lanceolate and broad-ovate leaves of Populus euphratica olive between Ejina Area in Inner Mongolia and Beijing Area[J]. Journal of MUC(Natural Sciences Edition),2016,25(3):85-91.
[14]杨树德,郑文菊,陈国仓,等.胡杨披针形叶与宽卵形叶的超微结构与光合特性的差异[J].西北植物学报,2005,25(1):14-21.Yang S D, Zheng W J, Chen G C, et al. Difference of ultrastructure and photosynthetic characteristics between lanceolate and broad-ovate leaves in Populus euphratica[J]. Acta Botanica Boreali-Occidentalia Sinica,2005,25(1):14-21.
[15]王海珍,韩路,徐雅丽,等.胡杨异形叶光合作用对光强与CO2浓度的响应[J].植物生态学报,2014,38(10):1099-1109.Wang H Z,Han L,Xu Y L,et al. Photosynthetic responses of the heteromorphic leaves in Populus euphratica to light intensity and CO2concentration[J]. Chinese Journal of Plant Ecology,2014,38(10):1099--1109.
[16]李雁玲,张肖,冯梅,等.胡杨(Populus euphratica)异形叶叶片内源激素特征研究[J].塔里木大学学报,2017,29(3):7-13.Li Y L,Zhang X,Fen M,et al. Characteristics of endhormones in leaf blade of Populus euphratica heteromorphic leaves[J]. Journal of Tarim University,2017,29(3):7--13.
[17]王玉丽,焦培培,顾亚亚,等.胡杨叶形变化与POD同工酶的关系研究[J].干旱区资源与环境,2013,27(4):181-186.Wang Y L,Jiao P P,Gu Y Y,et al. Study on the relationship between leaf shape change and POD enzyme of Populus euphratica[J]. Journal of Arid Land Resources and Environment,2013,27(4):181--186.
[18]秦少伟,李才林,郭忠忠,等.参与胡杨异形叶发生的发育相关基因表达模式及作用[J].基因组学与应用生物学,2017,36(11):4785--4792.Qin S W,Li C L,Guo Z Z,et al. Expression patterns and functions of genes related to the development of Populus euphratica Oliv. heteromorphic leaves morphogenesis[J]. Genomics and Applied Biology,2017,36(11):4785--4792.
[19]秦少伟,赵利峰,张娜,等. rot3、rot4、an和spk1在胡杨异形叶发生中的表达模式和作用[J].分子植物育种,2016,14(10):2630--2636.Qin S W,Zhao L F,Zhang N,et al. Expression patterns and effects of rot3, rot4, an and spk1 on the genesis of Populus euphratica Oliv. heteromorphic leaves[J]. Molecular Plant Breeding,2016,14(10):2630--2636.
[20]黄文娟,李志军,杨赵平,等.胡杨异形叶结构型性状及其与胸径关系[J].生态学杂志,2010,29(12):2347-2352.Huang W J, Li Z J, Yang Z P, et al. Heteromorphic leaf structural characteristics and their correlations with diameter at breast height of Populus euphratica[J]. Chinese Journal of Ecology,2010,29(12):2347-2352.
[21]岳宁,郑彩霞.胡杨异形叶蛋白质表达的电泳分析[J].安徽农业科学,2010,38(32):18058-18061.Yue N,Zheng C X. Electrophoresis analysis on the expressed proteinsin heteromorphic leaves of Populus euphratica Oliv.[J].Journal of Anhui Agri Sci,2010,38(32):18058-18061.
[22]苏培玺,张立新,杜明武,等.胡杨不同叶形光合特性、水分利用效率及其对加富CO2的响应[J].植物生态学报,2003,27(1):34--40.Su P X,Zhang L X,Du M W,et al. Photosynthetic character and water use efficiency of different leaf shapes of Populus euphratica and their response to CO2enrichment[J]. Acta Phytoecologica Sinica,2003,27(1):34--40.
[23]黄文娟,李志军,杨赵平,等.胡杨异形叶结构型性状及其相互关系[J].生态学报,2010,30(17):4636-4642.Huang W J,Li Z J,Yang Z P,et al. The structural traits of Populus euphratica heteromorphic leaves and their correlations[J]. Acta Ecologica Sinica,2010,30(17):4636--4642.
[24]刘帅飞,焦培培,李志军.灰叶胡杨异形叶的类型及其时空特征[J].干旱区研究,2016,33(5):1098--1103.Liu S F,Jiao P P,Li Z J. Diversifolious types and spatiotemporal characteristics of Populus pruinosa schrenk[J]. Arid Zone Research,2016,33(5):1098--1103.
[25] Weiner J. Allocation,plasticity and allometry in plants[J].Perspectives in Plant Ecology,Evolution and Systematics,2004,6(4):207--215.
[26] Zheng C, Qiu J, Jiang C, et al. Comparison of stomatal characteristics and photosynthesis of polymorphic Populus euphratica leaves[J]. Frontiers of Forestry in China,2007,2(1):87--93.
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