林下凋落物对土壤种子萌发的作用机制
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摘要
林下土壤种子库是森林植被的物种组成、种群延续和种群演替的物质基础。林下凋落物影响土壤种子库种子的萌发,幼苗的生长,进而影响到种群更新及森林群落的演替。凋落物对种子萌发的影响主要有三大作用机制,即物理作用机制化学作用机制、和生物作用机制。其中化学作用机制特别是化感作用对土壤种子库的影响尤其重要。本文将从这三个作用机制的层面,分析总结前人有关土壤种子库种子的萌发及其与林下凋落物、林下植被多样性的研究,论述土壤种子库研究面临的挑战,并对土壤种子库与生物多样性的研究进行了相关展望。
Soil seed bank of forest is the material basis of species composition,population growth and community succession for forest vegetation.Litter of forest understory affects seed germination,seedling growth,and population growth,thus affecting forest community succession.Litter affect soil seed bank through one of the three mechanisms,physical impact,chemical reaction,and biological interaction.Chemical reactions,especially allelopathy,play important roles in seed germination and seedling growth.In this paper,we analyzed and summarized previous studies on soil seed bank,litter and understory species diversity.We also discussed existing problems and future challenges on research of soil seed bank,litter and forest understory biodiversity.
Soil seed bank of forest is the material basis of species composition,population growth and community succession for forest vegetation.Litter of forest understory affects seed germination,seedling growth,and population growth,thus affecting forest community succession.Litter affect soil seed bank through one of the three mechanisms,physical impact,chemical reaction,and biological interaction.Chemical reactions,especially allelopathy,play important roles in seed germination and seedling growth.In this paper,we analyzed and summarized previous studies on soil seed bank,litter and understory species diversity.We also discussed existing problems and future challenges on research of soil seed bank,litter and forest understory biodiversity.
引文
[1]张玲,李广贺,张旭.土壤种子库研究综述[J].生态学杂志,2004,23(2):114-120.
[2]Kebromt,Tesfaye B.The role of soil seed banks in the rehabilitation of degraded hillslopes in Southern Wello Ethiopia[J].Biotropiea,2000,32(1):23-32.
[3]杨跃军,孙向阳,王保平.森林土壤种子库与天然更新[J].应用生态学报,2001,12(2):304-308.
[4]Bakker J P.Nature management by grazing and cutting[J].Journal of Applied Ecology,1990,5(4):128.
[5]于顺利,蒋高明.土壤种子库的研究进展及若干研究热点[J].植物生态学报,2003,27(4):552-560.
[6]安村青,林向阳,洪必恭.宝华山主要植被类型土壤种子库初探[J].植物生态学报,1996,20(1):41-50.
[7]陶大立,徐振邦,李昕.死、活地被物对红松伴生树种天然更新影响的实验研究[J].植物生态学报,1985,9(1):47-58.
[8]齐泽民,王开运,宋光煜,等.川西亚高山箭竹群落枯枝落叶层生物化学特性[J].生态学报,2004,24(6):1230-1236.
[9]张东来,毛子军,朱胜英,等.黑龙江省帽儿山林区6种主要林分类型凋落物研究[J].植物研究,2008,28(1):104-108.
[10]Persson S,Malmer N,Bo W.Leaf litter fall and soil acidity during half a century of secondary succession in a temperate deciduous forest[J].Vegetatio,1987,73(1):31-45.
[11]杨吉华,柳凯生.山丘地区森林保持水土效益的研究[J].水土保持学报,1993(3):47-52.
[12]郭伟,张健,黄玉梅,等.森林凋落物生态功能研究进展[J].安徽农业科学,2009,37(5):1984-1985.
[13]王俊,杨龙,李丹艳,等.凋落物覆盖及掩埋对不同质量的藜蒴种子萌发和幼苗生长的影响[J].生态环境学报,2008,17(5):1980-1985.
[14]Hitchmough J D.Urban landscape management.[J].Urban Landscape Management,1994.
[15]Gillman L N,Ogden J,Wright S D,et al.The influence of macro-itterfall and forest structure on litterfall damage to seedlings[J].Austral Ecology,2004,29(3):305-312.
[16]Pierson E A,Mack R N.The population biology of Bromus tectorum in forests:effect of disturbance,grazing,and litter on seedling establishment and reproduction[J].Oecologia,1990,84(4):526-533.
[17]刘尚华,石凤翎,吕世海,等.京西百花山区植物群落凋落物对土壤种子库的影响[J].水土保持通报,2008,28(2):41-47.
[18]周存宇.凋落物在森林生态系统中的作用及其研究进展[J].湖北农学院学报,2003,23(2):140-145.
[19]Manzoni S,Trofymow J A,Porporato J A.Stoichiometric controls on carbon,nitrogen,and phosphorus dynamics in decomposing litter[J].Ecological Monographs,2010,80(1):89-106.
[20]陈辉,唐明,刘安全,等.人工竹林凋落物中真菌群落的研究[J].西北林学院学报,1993(4):41-45.
[21]May F E,Ash J E.An Assessment of the Allelopathic Potential of Eucalyptus[J].Australian Journal of Botany,1990,38(3):245-254.
[22]Halvorson J J,Belesky D P,West M S.Inhibition of forage seed germination by leaf litter extracts of overstory hardwoods used in silvopastoral systems[J].Agroforestry Systems,2016,91(1):1-15.
[23]罗侠.天山云杉自毒作用的生化机制研究[D].乌鲁木齐:新疆农业大学,2006.
[24]李兆慧.天山云杉自毒作用与自毒物质的鉴定[D].杭州:浙江大学,2011.
[25]Reigosa M J,Gonzalez L.Forest ecosystems and allelopathy[M]//Allelopathy.Springer Netherlands,2006.
[26]Bu'Lock J D,Harbone J B.Phenolic compounds derived from shrikimate[J].1980.
[27]孔垂华.植物化感作用研究中应注意的问题[J].应用生态学报,1998,9(3):332-336.
[28]徐亮,包维楷,何永华.种子贮藏物质变化及其贮藏生理[J].种子,2003(5):60-63.
[29]杨期和,叶万辉,廖富林,等.植物化感物质对种子萌发的影响[J].生态学杂志,2005,24(12):1459-1465.
[30]陈慧玲,张新叶,杨彦伶.油茶水浸液对桔梗种子的化感效应[J].林业工程学报,2013,27(3):61-64.
[31]Jensen H A.Content of buried seeds in arable soil in Denmark and its relation to the weed population[J].Dansk Bot Arkiv,1969,41(4):314-20.
[32]Rice E L.Allelopathy[J].Encyclopedia of Entomology,1984(1):292-293.
[33]Muller W H.Volatile Materials Produced by Salvia leucophylla:Effects on Seedling Growth and Soil Bacteria[J].Botanical Gazette,1965,126(3):195-200.
[34]蒋有绪.川西亚高山冷杉林枯枝落叶层的群落学作用[J].植物生态学报,1981,5(2):4-13.
[35]李文华,赵景柱.生态学研究回顾与展望[M].北京:气象出版社,2004.
[36]盛海燕,常杰,殷现伟,等.濒危植物明党参种子散布和种子库动态研究[J].生物多样性,2002,10(3):269-273.
[37]Bewley,J.D.Seed Germination and Dormancy[J].The Plant Cell Online,1997,9(7):1055-1066.
[38]Dechoum M S,Castellani T T,Zalba S M,et al.Community structure,succession and invasibility in a seasonal deciduous forest in southern Brazil.[J].Biological Invasions,2015,17(6):1697-1712.
[39]Sevik,H.,Cetin,M.Effects of water stress on seed germination for select landscape plants[J].Pol.J.Environ.Stud,2015,24(2),689-693.
[40]Jimenezalfaro B,Silveira F O O,Fidelis A,et al.Seed germination traits can contribute better to plant community ecology[J].Journal of Vegetation Science,2016,27(3):637-645.
[41]张敏,朱教君,闫巧玲.光对种子萌发的影响机理研究进展[J].植物生态学报,2012,36(8):899-908.
[42]颜安,蒋平安,盛建东,等.玛纳斯河流域表层土壤盐分空间变异特征研究[J].土壤学报,2014(2):410-414.
[43]Zobel M,Kalamees R,Kersti Püssa,et al.Soil seed bank and vegetation in mixed coniferous forest stands with different disturbance regimes[J].Forest Ecology and Management,2007,250(1-2):0-76.
[44]Wagner M,Heinrich W,Jetschke G.Seed bank assembly in an unmanaged ruderal grassland recovering from long-term exposure to industrial emissions[J].Acta Oecologica,2006,30(3):342-352.
[45]Leck M A,Graveline K J.The seed bank of a freshwater tibal marsh[J].American Journal of Botany,1979,66(9):1006-1015.
[46]Peco B,Ortega M,Levassor C.Similarity between seed bank and vegetation in Mediterranean grassland:apredictive model[J].Journal of Vegetation Science,1998,9(6):815-828.
[47]Manzano P,Malo J E,et al.Sheep gut passage and survival of Mediterranean shrub seeds[J].Seed Science Research,2005,15(1):21-28.
[48]Henderson C B,Petersen K E,Redak RA.Spatial and temporal patterns in the bank and vetgetation of a desert grassland community[J].Journal of Ecology,1988,76(3):717-728.
[49]Roberts H A.Studies on the Weeds of Vegetable Crops:II.Effect of Six Years of Cropping on the Weed Seeds in the Soil[J].Journal of Ecology,1962,50(3):803-813.
[50]Roach D A.Buried seed and standing vegetation in two adjacent tundra habitats,northern Alaska[J].Oecologia,1983,60(3):359-364.
[51]刘明洪.长白山地区不同林型土壤种子库组成特征及与地上植被的关系[D].长春:东北师范大学,2010.
[52]Oswald A,Ransom J K.Striga control and improved farm productivity using crop rotation[J].Crop Protection,2001,20(2):113-120.
[53]Jensen H A.Content of buried seeds in arable soil in Denmark and its relation to the weed population[J].Dansk Bot Arkiv,1969,41(4):314-20.
[54]Thompson.Small-Scale Heterogeneity in the Seed Bank of An Acidic Grassland[J].Journal of Ecology,1986,74(3):733.
[55]Pickett S T A,Mcdonnell M J.CHAPTER 8–Seed Bank Dynamics in Temperate Deciduous Forest[J].Ecology of Soil Seed Banks,1989:123-147.
[56]尚占环,徐鹏彬,任国华,等.土壤种子库研究综述———植被系统中的作用及功能[J].草业学报,2009,18(2):175-183.
[2]Kebromt,Tesfaye B.The role of soil seed banks in the rehabilitation of degraded hillslopes in Southern Wello Ethiopia[J].Biotropiea,2000,32(1):23-32.
[3]杨跃军,孙向阳,王保平.森林土壤种子库与天然更新[J].应用生态学报,2001,12(2):304-308.
[4]Bakker J P.Nature management by grazing and cutting[J].Journal of Applied Ecology,1990,5(4):128.
[5]于顺利,蒋高明.土壤种子库的研究进展及若干研究热点[J].植物生态学报,2003,27(4):552-560.
[6]安村青,林向阳,洪必恭.宝华山主要植被类型土壤种子库初探[J].植物生态学报,1996,20(1):41-50.
[7]陶大立,徐振邦,李昕.死、活地被物对红松伴生树种天然更新影响的实验研究[J].植物生态学报,1985,9(1):47-58.
[8]齐泽民,王开运,宋光煜,等.川西亚高山箭竹群落枯枝落叶层生物化学特性[J].生态学报,2004,24(6):1230-1236.
[9]张东来,毛子军,朱胜英,等.黑龙江省帽儿山林区6种主要林分类型凋落物研究[J].植物研究,2008,28(1):104-108.
[10]Persson S,Malmer N,Bo W.Leaf litter fall and soil acidity during half a century of secondary succession in a temperate deciduous forest[J].Vegetatio,1987,73(1):31-45.
[11]杨吉华,柳凯生.山丘地区森林保持水土效益的研究[J].水土保持学报,1993(3):47-52.
[12]郭伟,张健,黄玉梅,等.森林凋落物生态功能研究进展[J].安徽农业科学,2009,37(5):1984-1985.
[13]王俊,杨龙,李丹艳,等.凋落物覆盖及掩埋对不同质量的藜蒴种子萌发和幼苗生长的影响[J].生态环境学报,2008,17(5):1980-1985.
[14]Hitchmough J D.Urban landscape management.[J].Urban Landscape Management,1994.
[15]Gillman L N,Ogden J,Wright S D,et al.The influence of macro-itterfall and forest structure on litterfall damage to seedlings[J].Austral Ecology,2004,29(3):305-312.
[16]Pierson E A,Mack R N.The population biology of Bromus tectorum in forests:effect of disturbance,grazing,and litter on seedling establishment and reproduction[J].Oecologia,1990,84(4):526-533.
[17]刘尚华,石凤翎,吕世海,等.京西百花山区植物群落凋落物对土壤种子库的影响[J].水土保持通报,2008,28(2):41-47.
[18]周存宇.凋落物在森林生态系统中的作用及其研究进展[J].湖北农学院学报,2003,23(2):140-145.
[19]Manzoni S,Trofymow J A,Porporato J A.Stoichiometric controls on carbon,nitrogen,and phosphorus dynamics in decomposing litter[J].Ecological Monographs,2010,80(1):89-106.
[20]陈辉,唐明,刘安全,等.人工竹林凋落物中真菌群落的研究[J].西北林学院学报,1993(4):41-45.
[21]May F E,Ash J E.An Assessment of the Allelopathic Potential of Eucalyptus[J].Australian Journal of Botany,1990,38(3):245-254.
[22]Halvorson J J,Belesky D P,West M S.Inhibition of forage seed germination by leaf litter extracts of overstory hardwoods used in silvopastoral systems[J].Agroforestry Systems,2016,91(1):1-15.
[23]罗侠.天山云杉自毒作用的生化机制研究[D].乌鲁木齐:新疆农业大学,2006.
[24]李兆慧.天山云杉自毒作用与自毒物质的鉴定[D].杭州:浙江大学,2011.
[25]Reigosa M J,Gonzalez L.Forest ecosystems and allelopathy[M]//Allelopathy.Springer Netherlands,2006.
[26]Bu'Lock J D,Harbone J B.Phenolic compounds derived from shrikimate[J].1980.
[27]孔垂华.植物化感作用研究中应注意的问题[J].应用生态学报,1998,9(3):332-336.
[28]徐亮,包维楷,何永华.种子贮藏物质变化及其贮藏生理[J].种子,2003(5):60-63.
[29]杨期和,叶万辉,廖富林,等.植物化感物质对种子萌发的影响[J].生态学杂志,2005,24(12):1459-1465.
[30]陈慧玲,张新叶,杨彦伶.油茶水浸液对桔梗种子的化感效应[J].林业工程学报,2013,27(3):61-64.
[31]Jensen H A.Content of buried seeds in arable soil in Denmark and its relation to the weed population[J].Dansk Bot Arkiv,1969,41(4):314-20.
[32]Rice E L.Allelopathy[J].Encyclopedia of Entomology,1984(1):292-293.
[33]Muller W H.Volatile Materials Produced by Salvia leucophylla:Effects on Seedling Growth and Soil Bacteria[J].Botanical Gazette,1965,126(3):195-200.
[34]蒋有绪.川西亚高山冷杉林枯枝落叶层的群落学作用[J].植物生态学报,1981,5(2):4-13.
[35]李文华,赵景柱.生态学研究回顾与展望[M].北京:气象出版社,2004.
[36]盛海燕,常杰,殷现伟,等.濒危植物明党参种子散布和种子库动态研究[J].生物多样性,2002,10(3):269-273.
[37]Bewley,J.D.Seed Germination and Dormancy[J].The Plant Cell Online,1997,9(7):1055-1066.
[38]Dechoum M S,Castellani T T,Zalba S M,et al.Community structure,succession and invasibility in a seasonal deciduous forest in southern Brazil.[J].Biological Invasions,2015,17(6):1697-1712.
[39]Sevik,H.,Cetin,M.Effects of water stress on seed germination for select landscape plants[J].Pol.J.Environ.Stud,2015,24(2),689-693.
[40]Jimenezalfaro B,Silveira F O O,Fidelis A,et al.Seed germination traits can contribute better to plant community ecology[J].Journal of Vegetation Science,2016,27(3):637-645.
[41]张敏,朱教君,闫巧玲.光对种子萌发的影响机理研究进展[J].植物生态学报,2012,36(8):899-908.
[42]颜安,蒋平安,盛建东,等.玛纳斯河流域表层土壤盐分空间变异特征研究[J].土壤学报,2014(2):410-414.
[43]Zobel M,Kalamees R,Kersti Püssa,et al.Soil seed bank and vegetation in mixed coniferous forest stands with different disturbance regimes[J].Forest Ecology and Management,2007,250(1-2):0-76.
[44]Wagner M,Heinrich W,Jetschke G.Seed bank assembly in an unmanaged ruderal grassland recovering from long-term exposure to industrial emissions[J].Acta Oecologica,2006,30(3):342-352.
[45]Leck M A,Graveline K J.The seed bank of a freshwater tibal marsh[J].American Journal of Botany,1979,66(9):1006-1015.
[46]Peco B,Ortega M,Levassor C.Similarity between seed bank and vegetation in Mediterranean grassland:apredictive model[J].Journal of Vegetation Science,1998,9(6):815-828.
[47]Manzano P,Malo J E,et al.Sheep gut passage and survival of Mediterranean shrub seeds[J].Seed Science Research,2005,15(1):21-28.
[48]Henderson C B,Petersen K E,Redak RA.Spatial and temporal patterns in the bank and vetgetation of a desert grassland community[J].Journal of Ecology,1988,76(3):717-728.
[49]Roberts H A.Studies on the Weeds of Vegetable Crops:II.Effect of Six Years of Cropping on the Weed Seeds in the Soil[J].Journal of Ecology,1962,50(3):803-813.
[50]Roach D A.Buried seed and standing vegetation in two adjacent tundra habitats,northern Alaska[J].Oecologia,1983,60(3):359-364.
[51]刘明洪.长白山地区不同林型土壤种子库组成特征及与地上植被的关系[D].长春:东北师范大学,2010.
[52]Oswald A,Ransom J K.Striga control and improved farm productivity using crop rotation[J].Crop Protection,2001,20(2):113-120.
[53]Jensen H A.Content of buried seeds in arable soil in Denmark and its relation to the weed population[J].Dansk Bot Arkiv,1969,41(4):314-20.
[54]Thompson.Small-Scale Heterogeneity in the Seed Bank of An Acidic Grassland[J].Journal of Ecology,1986,74(3):733.
[55]Pickett S T A,Mcdonnell M J.CHAPTER 8–Seed Bank Dynamics in Temperate Deciduous Forest[J].Ecology of Soil Seed Banks,1989:123-147.
[56]尚占环,徐鹏彬,任国华,等.土壤种子库研究综述———植被系统中的作用及功能[J].草业学报,2009,18(2):175-183.