异质性水分环境中克隆整合对活血丹生物量分配及叶片结构特征的影响
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摘要
喀斯特石漠化环境有着高度的生境异质性,异质性生境中土被不连续,土壤瘠薄,岩溶漏斗上的土壤保水性差,严重制约着喀斯特植被的生长及分布。为探究克隆植物在喀斯特地区的适应策略,本研究以喀斯特黄色石灰土为基质,选用克隆植物活血丹(Glechoma longituba),以一个节间连接的两个分株为材料,保持节间连接或切断,种植于相邻花盆中,并施以不同浇水量,以明确不同水分可用性水平下克隆整合对活血丹生物量积累、生物量分配、叶片气孔及叶片组织特征的影响。结果显示,克隆整合显著促进活血丹生物量的积累及对根、叶的生物量分配;增加了活血丹叶气孔导度,降低了气孔指数;叶海绵组织受克隆整合影响较小,但栅栏组织及栅海比(栅栏组织/海绵组织)表现为非整合分株高于整合分株。本研究表明,克隆整合可增加活血丹胁迫分株对根、叶的投资,并以更佳的叶气孔、组织适应策略提高其在喀斯特生境中的生存与适应。
Because of discontinuous soil coverage, poor soil fertility and soil water scarcity, the high heterogeneity of karst habitats seriously restricts the growth and distribution of local plants. To clarify the adaptive strategies of clonal plants in karst area,two connected ramets with a single internode of Glechoma longituba were planted in yellow calcareous soil. Then,we cut off the half of the internode and made the ramets disconnected,supplied different amounts of water to clarify the effects of the clonal integration on biomass accumulation,distribution,stomata and tissue characteristics of leaves under different treatments. The results showed that clonal integration significantly promoted the biomass accumulation of G. longituba,and increased biomass allocation to roots and leaves. The stomatal conductance were increased and the stomatal index were decreased by clonal integration of leaves. There was less affected on leaves sponge tissue,but the palisade tissue and the ratio of palisade to spongy tissue showed that the non-clonal integration ramets were higher than the integrated ramets. The clonal integration could increase the investment of roots and leaves of G. longituba,and further promoted clonal plant survival and adaptation in karst habitats with better leaves stomata and tissue structure.
Because of discontinuous soil coverage, poor soil fertility and soil water scarcity, the high heterogeneity of karst habitats seriously restricts the growth and distribution of local plants. To clarify the adaptive strategies of clonal plants in karst area,two connected ramets with a single internode of Glechoma longituba were planted in yellow calcareous soil. Then,we cut off the half of the internode and made the ramets disconnected,supplied different amounts of water to clarify the effects of the clonal integration on biomass accumulation,distribution,stomata and tissue characteristics of leaves under different treatments. The results showed that clonal integration significantly promoted the biomass accumulation of G. longituba,and increased biomass allocation to roots and leaves. The stomatal conductance were increased and the stomatal index were decreased by clonal integration of leaves. There was less affected on leaves sponge tissue,but the palisade tissue and the ratio of palisade to spongy tissue showed that the non-clonal integration ramets were higher than the integrated ramets. The clonal integration could increase the investment of roots and leaves of G. longituba,and further promoted clonal plant survival and adaptation in karst habitats with better leaves stomata and tissue structure.
引文
1.Hutchings M J,John E A.The effects of environmental heterogeneity on root growth and root/shoot partitioning[J].Annals of Botany,2004,94(1):1-8.
2.李阳兵.西南岩溶山地生态环境研究进展与若干问题[J].云南地理环境研究,2006,18(2):61-65.Li Y B.Some progresses and problems of research on ecological reconstruction of karst mountain areas in Southwest China[J].Yunnan Geographic Environment Research,2006,18(2):61-65.
3.刘方,王世杰,刘元生,等.喀斯特石漠化过程土壤质量变化及生态环境影响评价[J].生态学报,2005,25(3):639-644.Liu F,Wang S J,Liu Y S,et al.Changes of soil quality in the process of karst rocky desertification and evaluation of impact on ecological environment[J].Acta Ecologica Sinica,2005,25(3):639-644.
4.张伟,陈洪松,王克林,等.喀斯特地区典型峰丛洼地旱季表层土壤水分空间变异性初探[J].土壤学报,2006,43(4):554-562.Zhang W,Chen H S,Wang K L,et al.Spatial variability of surface soil water in typical depressions between hills in karst region in dry season[J].Acta Pedologica Sinica,2006,43(4):554-562.
5.Li Y,Chen J S,Xue G,et al.Effect of clonal integration on nitrogen cycling in rhizosphere of rhizomatous clonal plant,Phyllostachys bissetii,under heterogeneous light[J].Science of the Total Environment,2018,628-629:594-602.
6.Stuefer J F.Potential and limitations of current concepts regarding the response of clonal plants to environmental heterogeneity[J].Vegetatio,1996,127(1):55-70.
7.何军,赵聪蛟,清华,等.土壤水分条件对克隆植物互花米草表型可塑性的影响[J].生态学报,2009,29(7):3518-3524.He J,Zhao C J,Qing H,et al.Effect of soil-water condition on morphological plasticity of clonal plant Spartina alterniflora[J].Acta Ecologica Sinica,2009,29(7):3518-3524.
8.Meng L S.Compound synthesis or growth and development of roots/stomata regulate plant drought tolerance or water use efficiency/water uptake efficiency[J].Journal of Agricultural and Food Chemistry,2018,66(14):3595-3604.
9.Cortés A J,Monserrate F A,Ramírez-Villegas J,et al.Drought tolerance in wild plant populations:the case of common beans(Phaseolus vulgaris L.)[J].PLo S One,2013,8(5):e62898.
10.孙三杰,李建明,宗建伟,等.亚低温与干旱胁迫对番茄幼苗根系形态及叶片结构的影响[J].应用生态学报,2012,23(11):3027-3032.Sun S J,Li J M,Zong J W,et al.Effects of sub-low temperature and drought stress on root morphology and leaf structure of tomato seedlings[J].Chinese Journal of Applied E-cology,2012,23(11):3027-3032.
11.胡忠良,潘根兴,李恋卿,等.贵州喀斯特山区不同植被下土壤C、N、P含量和空间异质性[J].生态学报,2009,29(8):4187-4195.Hu Z L,Pan G X,Li L Q,et al.Changes in pools and heterogeneity of soil organic carbon,nitrogen and phosphorus under different vegetation types in karst mountainous area of central Guizhou Province,China[J].Acta Ecologica Sinica,2009,29(8):4187-4195.
12.刘方,王世杰,罗海波,等.喀斯特森林生态系统的小生境及其土壤异质性[J].土壤学报,2008,45(6):1055-1062.Liu F,Wang S J,Luo H B,et al.Micro-habitats in karst forest ecosystem and variability of soil[J].Acta Pedologica Sinica,2008,45(6):1055-1062.
13.Stueffer J F,De Kroon H,During H J.Exploitation of environmental hetergeneity by spatial division of labor in a clonal plant[J].Functional Ecology,1996,10(3):328-334.
14.董鸣.异质性生境中的植物克隆生长:风险分摊[J].植物生态学报,1996,20(6):543-548.Dong M.Plant clonal growth in heterogeneous habitats:Risk-spreading[J].Acta Phytoecologica Sinica,1996,20(6):543-548.
15.董鸣.资源异质性环境中的植物克隆生长:觅食行为[J].植物学报,1996,38(10):828-835.Dong M.Clonal growth in plants in relation to resource heterogeneity:Foraging behavior[J].Acta Botanica Sinica,1996,38(10):828-835.
16.De Kroons H,Hutchings M J.Morphological plasticity in clonal plants:the foraging concept reconsidered[J].Journal of Ecology,1995,83(1):143-152.
17.Pan J J,Price J S.Fitness and evolution in clonal plants:the impact of clonal growth[J].Evolutionary Ecology,2001,15(4-6):583-600.
18.Van Der Hoeven E C,De Kroon H,During H J.Fine-scale spatial distribution of leaves and shoots of two chalk grassland perennials[J].Vegetatio,1990,86(2):151-160.
19.王俊峰,冯玉龙.光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响[J].植物生态学报,2004,28(6):781-786.Wang J F,Feng Y L.The effect of light intensity on biomass allocation,leaf morphology and relative growth rate of two invasive plants[J].Acta Phytoecologica Sinica,2004,28(6):781-786.
20.徐高峰,申时才,张付斗,等.土壤水分对薇甘菊不同繁殖体单位存活能力和植株表型可塑性影响[J].中国农业科学,2013,46(15):3134-3141.Xu G F,Shen S C,Zhang F D,et al.Effect of soil-water conditions on survival rate and morphological plasticity of clonal plant Mikania micrantha H.B.Kunth[J].Scientia Agricultura Sinica,2013,46(15):3134-3141.
21.武晓静.资源斑块交互性与一致性对根茎克隆植物生长特性与克隆整合的影响[D].武汉:华中农业大学,2015.Wu X J.Effects of reciprocal patchiness and coincident patchiness of resources on the cloning features and clonal integration of Iris japonica Thunb.[D].Wuhan:Huazhong Agricultural University,2015.
22.钱永强.异质性水分胁迫下野牛草克隆分株间生理整合及其调控机理[D].北京:中国林业科学研究院,2009.Qian Y Q.Physiological integration and its regulation between Inter-ramet of Buffalograss under heterogeneous water stress[D].Beijing:Chinese Academy of Forestry,2009.
23.Roiloa S R,Rodríguez-Echeverría S,De La Pe1a E,et al.Physiological integration increases the survival and growth of the clonal invader Carpobrotus edulis[J].Biological Invasions,2010,12(6):1815-1823.
24.吴志庄,杨志和,厉月桥,等.木本能源植物黄连木苗期生物量分配规律与构成因素分析[J].西北林学院学报,2015,30(6):28-32.Wu Z Z,Yang Z H,Li Y Q,et al.Comprehensive analysis on the distribution pattern and constituent factors affecting seedling biomass of woody energy plant Pistacia chinensis[J].Journal of Northwest Forestry University,2015,30(6):28-32.
25.Zhou J,Dong B C,Alpert P,et al.Effects of soil nutrient heterogeneity on intraspecific competition in the invasive,clonal plant Alternanthera philoxeroides[J].Annals of Botany,2012,109(4):813-818.
26.董鸣,张淑敏,陈玉福.匍匐茎草本蛇莓对基质养分条件的克隆可塑性[J].植物学报,2000,42(5):518-522.Dong M,Zhang S M,Chen Y F.Clonal plasticity in response to nutrient availability in the stoloniferous herb,Duchesnea indica[J].Acta Botanica Sinica,2000,42(5):518-522.
27.Thorpe M R,Warrit B,Landsberg J J.Responses of apple leaf stomata:a model for single leaves and a whole tree[J].Plant,Cell&Environment,1980,3(1):23-27.
28.Kubota F,Yoshimura Y,Agata W.Stomatal movement and CO2exchange rate of sweet potato plant(Ipomoea batatas Lam.)in relation to water environments:a comparison between native and improved varieties[J].Journal of the Faculty of Agriculture,Kyushu University,1993,38(1-2):97-110.
29.Wang Y,Chen X,Xiang C B.Stomatal density and bio-water saving[J].Journal of Integrative Plant Biology,2007,49(10):1435-1444.
30.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.
31.孟庆杰,王光全,董绍锋,等.桃叶片组织解剖结构特征与其抗旱性关系的研究[J].干旱地区农业研究,2004,22(3):123-126.Meng Q J,Wang G Q,Dong S F,et al.Relation between leaf tissue parameters and drought resistance of peaches[J].Agricultural Research in the Arid Areas,2004,22(3):123-126.
32.常英俏,徐文远,穆立蔷,等.干旱胁迫对3种观赏灌木叶片解剖结构的影响及抗旱性分析[J].东北林业大学学报,2012,40(3):36-40.Chang Y Q,Xu W Y,Mu L H,et al.Effects of drought stress on anatomical structure of leaves of three species of shrubs and their drought resistances[J].Journal of Northeast Forestry University,2012,40(3):36-40.
33.陈国菊,杨暹,吴筱颖.干旱胁迫对花椰菜叶片细胞保护酶及超微结构的影响[J].中国蔬菜,2002(2):8-11.Chen G J,Yang X,Wu X Y.The effects of drought stress on cell-protecting enzymes and ultrastructure of cauliflower leaf cells[J].Chinese Vegetables,2002(2):8-11.
2.李阳兵.西南岩溶山地生态环境研究进展与若干问题[J].云南地理环境研究,2006,18(2):61-65.Li Y B.Some progresses and problems of research on ecological reconstruction of karst mountain areas in Southwest China[J].Yunnan Geographic Environment Research,2006,18(2):61-65.
3.刘方,王世杰,刘元生,等.喀斯特石漠化过程土壤质量变化及生态环境影响评价[J].生态学报,2005,25(3):639-644.Liu F,Wang S J,Liu Y S,et al.Changes of soil quality in the process of karst rocky desertification and evaluation of impact on ecological environment[J].Acta Ecologica Sinica,2005,25(3):639-644.
4.张伟,陈洪松,王克林,等.喀斯特地区典型峰丛洼地旱季表层土壤水分空间变异性初探[J].土壤学报,2006,43(4):554-562.Zhang W,Chen H S,Wang K L,et al.Spatial variability of surface soil water in typical depressions between hills in karst region in dry season[J].Acta Pedologica Sinica,2006,43(4):554-562.
5.Li Y,Chen J S,Xue G,et al.Effect of clonal integration on nitrogen cycling in rhizosphere of rhizomatous clonal plant,Phyllostachys bissetii,under heterogeneous light[J].Science of the Total Environment,2018,628-629:594-602.
6.Stuefer J F.Potential and limitations of current concepts regarding the response of clonal plants to environmental heterogeneity[J].Vegetatio,1996,127(1):55-70.
7.何军,赵聪蛟,清华,等.土壤水分条件对克隆植物互花米草表型可塑性的影响[J].生态学报,2009,29(7):3518-3524.He J,Zhao C J,Qing H,et al.Effect of soil-water condition on morphological plasticity of clonal plant Spartina alterniflora[J].Acta Ecologica Sinica,2009,29(7):3518-3524.
8.Meng L S.Compound synthesis or growth and development of roots/stomata regulate plant drought tolerance or water use efficiency/water uptake efficiency[J].Journal of Agricultural and Food Chemistry,2018,66(14):3595-3604.
9.Cortés A J,Monserrate F A,Ramírez-Villegas J,et al.Drought tolerance in wild plant populations:the case of common beans(Phaseolus vulgaris L.)[J].PLo S One,2013,8(5):e62898.
10.孙三杰,李建明,宗建伟,等.亚低温与干旱胁迫对番茄幼苗根系形态及叶片结构的影响[J].应用生态学报,2012,23(11):3027-3032.Sun S J,Li J M,Zong J W,et al.Effects of sub-low temperature and drought stress on root morphology and leaf structure of tomato seedlings[J].Chinese Journal of Applied E-cology,2012,23(11):3027-3032.
11.胡忠良,潘根兴,李恋卿,等.贵州喀斯特山区不同植被下土壤C、N、P含量和空间异质性[J].生态学报,2009,29(8):4187-4195.Hu Z L,Pan G X,Li L Q,et al.Changes in pools and heterogeneity of soil organic carbon,nitrogen and phosphorus under different vegetation types in karst mountainous area of central Guizhou Province,China[J].Acta Ecologica Sinica,2009,29(8):4187-4195.
12.刘方,王世杰,罗海波,等.喀斯特森林生态系统的小生境及其土壤异质性[J].土壤学报,2008,45(6):1055-1062.Liu F,Wang S J,Luo H B,et al.Micro-habitats in karst forest ecosystem and variability of soil[J].Acta Pedologica Sinica,2008,45(6):1055-1062.
13.Stueffer J F,De Kroon H,During H J.Exploitation of environmental hetergeneity by spatial division of labor in a clonal plant[J].Functional Ecology,1996,10(3):328-334.
14.董鸣.异质性生境中的植物克隆生长:风险分摊[J].植物生态学报,1996,20(6):543-548.Dong M.Plant clonal growth in heterogeneous habitats:Risk-spreading[J].Acta Phytoecologica Sinica,1996,20(6):543-548.
15.董鸣.资源异质性环境中的植物克隆生长:觅食行为[J].植物学报,1996,38(10):828-835.Dong M.Clonal growth in plants in relation to resource heterogeneity:Foraging behavior[J].Acta Botanica Sinica,1996,38(10):828-835.
16.De Kroons H,Hutchings M J.Morphological plasticity in clonal plants:the foraging concept reconsidered[J].Journal of Ecology,1995,83(1):143-152.
17.Pan J J,Price J S.Fitness and evolution in clonal plants:the impact of clonal growth[J].Evolutionary Ecology,2001,15(4-6):583-600.
18.Van Der Hoeven E C,De Kroon H,During H J.Fine-scale spatial distribution of leaves and shoots of two chalk grassland perennials[J].Vegetatio,1990,86(2):151-160.
19.王俊峰,冯玉龙.光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响[J].植物生态学报,2004,28(6):781-786.Wang J F,Feng Y L.The effect of light intensity on biomass allocation,leaf morphology and relative growth rate of two invasive plants[J].Acta Phytoecologica Sinica,2004,28(6):781-786.
20.徐高峰,申时才,张付斗,等.土壤水分对薇甘菊不同繁殖体单位存活能力和植株表型可塑性影响[J].中国农业科学,2013,46(15):3134-3141.Xu G F,Shen S C,Zhang F D,et al.Effect of soil-water conditions on survival rate and morphological plasticity of clonal plant Mikania micrantha H.B.Kunth[J].Scientia Agricultura Sinica,2013,46(15):3134-3141.
21.武晓静.资源斑块交互性与一致性对根茎克隆植物生长特性与克隆整合的影响[D].武汉:华中农业大学,2015.Wu X J.Effects of reciprocal patchiness and coincident patchiness of resources on the cloning features and clonal integration of Iris japonica Thunb.[D].Wuhan:Huazhong Agricultural University,2015.
22.钱永强.异质性水分胁迫下野牛草克隆分株间生理整合及其调控机理[D].北京:中国林业科学研究院,2009.Qian Y Q.Physiological integration and its regulation between Inter-ramet of Buffalograss under heterogeneous water stress[D].Beijing:Chinese Academy of Forestry,2009.
23.Roiloa S R,Rodríguez-Echeverría S,De La Pe1a E,et al.Physiological integration increases the survival and growth of the clonal invader Carpobrotus edulis[J].Biological Invasions,2010,12(6):1815-1823.
24.吴志庄,杨志和,厉月桥,等.木本能源植物黄连木苗期生物量分配规律与构成因素分析[J].西北林学院学报,2015,30(6):28-32.Wu Z Z,Yang Z H,Li Y Q,et al.Comprehensive analysis on the distribution pattern and constituent factors affecting seedling biomass of woody energy plant Pistacia chinensis[J].Journal of Northwest Forestry University,2015,30(6):28-32.
25.Zhou J,Dong B C,Alpert P,et al.Effects of soil nutrient heterogeneity on intraspecific competition in the invasive,clonal plant Alternanthera philoxeroides[J].Annals of Botany,2012,109(4):813-818.
26.董鸣,张淑敏,陈玉福.匍匐茎草本蛇莓对基质养分条件的克隆可塑性[J].植物学报,2000,42(5):518-522.Dong M,Zhang S M,Chen Y F.Clonal plasticity in response to nutrient availability in the stoloniferous herb,Duchesnea indica[J].Acta Botanica Sinica,2000,42(5):518-522.
27.Thorpe M R,Warrit B,Landsberg J J.Responses of apple leaf stomata:a model for single leaves and a whole tree[J].Plant,Cell&Environment,1980,3(1):23-27.
28.Kubota F,Yoshimura Y,Agata W.Stomatal movement and CO2exchange rate of sweet potato plant(Ipomoea batatas Lam.)in relation to water environments:a comparison between native and improved varieties[J].Journal of the Faculty of Agriculture,Kyushu University,1993,38(1-2):97-110.
29.Wang Y,Chen X,Xiang C B.Stomatal density and bio-water saving[J].Journal of Integrative Plant Biology,2007,49(10):1435-1444.
30.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.
31.孟庆杰,王光全,董绍锋,等.桃叶片组织解剖结构特征与其抗旱性关系的研究[J].干旱地区农业研究,2004,22(3):123-126.Meng Q J,Wang G Q,Dong S F,et al.Relation between leaf tissue parameters and drought resistance of peaches[J].Agricultural Research in the Arid Areas,2004,22(3):123-126.
32.常英俏,徐文远,穆立蔷,等.干旱胁迫对3种观赏灌木叶片解剖结构的影响及抗旱性分析[J].东北林业大学学报,2012,40(3):36-40.Chang Y Q,Xu W Y,Mu L H,et al.Effects of drought stress on anatomical structure of leaves of three species of shrubs and their drought resistances[J].Journal of Northeast Forestry University,2012,40(3):36-40.
33.陈国菊,杨暹,吴筱颖.干旱胁迫对花椰菜叶片细胞保护酶及超微结构的影响[J].中国蔬菜,2002(2):8-11.Chen G J,Yang X,Wu X Y.The effects of drought stress on cell-protecting enzymes and ultrastructure of cauliflower leaf cells[J].Chinese Vegetables,2002(2):8-11.
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