杉木种子萌发及幼苗生长对光强的响应
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摘要
设置不同光强梯度(透光率分别为100%、40%、20%、10%和5%,光照强度PPFD分别为201.3、77.0、37.5、19.2、9.8μmol·m~(-2)·s~(-1)),研究光对杉木种子萌发和幼苗早期生长的影响,分析杉木种子萌发、幼苗存活、生长、形态响应、生物量积累及其分配格局对不同光环境的响应策略.结果表明:杉木种子的萌发率、存活率、建植率和萌发指数在不同光强梯度下均有显著差异,且40%的透光率是种子最适萌发条件,萌发率最高,而全光照下存活率和建植率最高;随光照强度的减弱,杉木幼苗茎长增大,根长、子叶长、子叶厚、真叶数呈降低趋势,而基径在各光照强度间无显著差异;总生物量、根生物量、茎生物量、叶生物量均表现为全光照下最大.随着光照强度的减弱,光合组织与非光合组织生物量比、叶生物量比呈降低趋势,茎生物量比呈增加趋势,根冠比和根生物量比无显著差异.弱光环境促进杉木种子萌发,不利于杉木幼苗存活和生长.在弱光环境下,杉木幼苗通过增大茎生物量来提高对弱光环境的耐受力.
The effect of light intensity on the seed germination and seedling growth of Chinese fir under different light intensities( 100%,40%,20%,10%,5% of full light,and the PPFD was201.3,77.0,37.5,19.2,9.8 μmol·m~(-2)·s~(-1),respectively) was investigated,and the adaptive strategy of seed germination,seedling survival,growth,morphological plasticity,biomass accumulation and allocation under different light intensities was explored in this paper. The results showed that light intensity significantly affected the germination rate,survival rate,establishment rate and germination index. Germination rate reached the maximum under 40% light intensity,while survival rate and establishment rate reached the maximum at 100% light intensity. With the light intensity decreased,the stem length increased,while the root length,cotyledon length,cotyledon thickness and euphylla number declined,and basal stem diameter had no significant difference among different light intensities. The total biomass,root biomass,stem biomass and leaf biomass were the highest under 100% light intensity. With the light intensity decreased,the photosynthesis non-photosynthesis biomass ratio and leaf biomass ratio declined,while stem biomass ratio increased,the root to shoot ratio and root biomass ratio had no significant difference among different light intensities. Low light promoted seed germination,but seedlings grew slowly with high mortality under low light. The accumulation of biomass in stem increased the plant tolerance to low light.
The effect of light intensity on the seed germination and seedling growth of Chinese fir under different light intensities( 100%,40%,20%,10%,5% of full light,and the PPFD was201.3,77.0,37.5,19.2,9.8 μmol·m~(-2)·s~(-1),respectively) was investigated,and the adaptive strategy of seed germination,seedling survival,growth,morphological plasticity,biomass accumulation and allocation under different light intensities was explored in this paper. The results showed that light intensity significantly affected the germination rate,survival rate,establishment rate and germination index. Germination rate reached the maximum under 40% light intensity,while survival rate and establishment rate reached the maximum at 100% light intensity. With the light intensity decreased,the stem length increased,while the root length,cotyledon length,cotyledon thickness and euphylla number declined,and basal stem diameter had no significant difference among different light intensities. The total biomass,root biomass,stem biomass and leaf biomass were the highest under 100% light intensity. With the light intensity decreased,the photosynthesis non-photosynthesis biomass ratio and leaf biomass ratio declined,while stem biomass ratio increased,the root to shoot ratio and root biomass ratio had no significant difference among different light intensities. Low light promoted seed germination,but seedlings grew slowly with high mortality under low light. The accumulation of biomass in stem increased the plant tolerance to low light.
引文
[1]Chen S-B(陈圣宾),Song A-Q(宋爱琴),Li Z-J(李振基).Research advance in response of forest seedling regeneration to light environmental heterogeneity.Chinese Journal of Applied Ecology(应用生态学报),2005,16(2):365-370(in Chinese)
[2]Du XJ,Guo QF,Gao XM,et al.Seed rain,soil seed bank,seed loss and regeneration of Castanopsis fargesii(Fagaceae)in a subtropical evergreen broad-leaved forest.Forest Ecology and Management,2007,238:212-219
[3]Scholes J,Press M,Zipperlen S.Differences in light energy utilisation and dissipation between dipterocarp rain forest tree seedlings.Oecologia,1996,109:41-48
[4]Han Y-Z(韩有志),Wang Z-Q(王政权).Spatial heterogeneity and forest regeneration.Chinese Journal of Applied Ecology(应用生态学报),2002,13(5):615-619(in Chinese)
[5]Nicotra AB,Chazdon RL,Iriarte SVB.Spatial heterogeneity of light and woody seedling regeneration in tropical wet forests.Ecology,1999,80:1908-1926
[6]Diaci J.Regeneration dynamics in a Norway spruce plantation on a silver fir-beech forest site in the Slovenian Alps.Forest Ecology and Management,2002,161:27-38
[7]Lai J-S(赖江山),Li Q-M(李庆梅),Xie Z-Q(谢宗强).Seed germinating characteristics of the endangered plant Abies chensiensis.Chinese Journal of Plant Ecology(植物生态学报),2003,27(5):661-666(in Chinese)
[8]Wu Y(吴彦),Liu Q(刘庆),He H(何海),et al.Effects of light and temperature on seed germination of Picea asperata and Betula albo-sinensis.Chinese Journal of Applied Ecology(应用生态学报),2004,15(12):2229-2232(in Chinese)
[9]Yan X-F(闫兴富),Cao M(曹敏).Effects of light intensity on seed germination and seedling early growth of Shorea wantianshuea.Chinese Journal of Applied Ecology(应用生态学报),2007,18(1):23-29(in Chinese)
[10]Benigno GR,Mulualem T,Guillermo,et al.Seed germination and seedling establishment of Neotropical dry forest species in response to temperature and light conditions.Journal of Forestry Research,2009,20:99-104
[11]Mc Kinnon LM,Mitchell AK.Photoprotection,not increased growth,characterizes the response of Engelmann spruce(Picea engelmannii)seedlings to high light,even when resources are plentiful.New Phytologist,2003,160:69-79
[12]Mc Connaughay KDM,Coleman JS.Biomass allocation in plants:Ontogeny or optimality?A test along three resource gradients.Ecology,1999,80:2581-2593
[13]Grubb PJ,Lee WG,Kollmann J,et al.Interaction of irradiance and soil nutrient supply on growth of seedlings of ten European tall-shrub species and Fagus sylvatica.Journal of Ecology,1996,84:827-840
[14]Evans JR,Poorter H.Photosynthetic acclimation of plants to growth irradiance:The relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain.Plant,Cell&Environment,2001,24:755-767
[15]Mokany K,Raison R,Prokushkin AS.Critical analysis of root:Shoot ratios in terrestrial biomes.Global Change Biology,2006,12:84-96
[16]Lockhart BR,Gardiner ES,Hodges JD,et al.Carbon allocation and morphology of cherrybark oak seedlings and sprouts under three light regimes.Annals of Forest Science,2008,65:801
[17]Stearns SC.The Evolution of Life Histories.Oxford:Oxford University Press,1992
[18]Bloom AJ,Chapin FS,Mooney HA.Resource limitation in plants:An economic analogy.Annual Review of Ecology and Systematics,1985,16:363-392
[19]Geng H-L(耿浩林),Wang Y-H(王玉辉),Wang FY(王风玉),et al.The dynamics of root-shoot ratio and its environmental effective factors of recovering Leymus chinensis steppe vegetation in Inner Mongolia.Acta Ecologica Sinica(生态学报),2008,28(10):4629-4634(in Chinese)
[20]Zhang M(张敏).The Germination,Seedling Survival and Early Growth of the Main Tree Species in Response to Light Regimes in Secondary Forests,Northeast China.Ph D Thesis.Shenyang:Institute Appled Ecology,Chinese Academy of Sciences,2012(in Chinese)
[21]Portsmuth A,Niinemets.Structural and physiological plasticity in response to light and nutrients in five temperate deciduous woody species of contrasting shade tolerance.Functional Ecology,2007,21:61-77
[22]Yu X-T(俞新妥).A summary of the studies on Chinese fir in 2000-2005.Ⅰ.The research development on physiological ecology of Chinese fir.Journal of Fujian College of Forestry(福建林学院学报),2006,26(2):177-185(in Chinese)
[23]Yu X-T(俞新妥).Summary of the studies on the plantation productivity and nutrient cycling in Chinese fir plantation ecosystem.World Forestry Research(世界林业研究),1993,2(3):80-86(in Chinese)
[24]Chen C-Y(陈楚莹),Zhang J-W(张家武),Zhou C-L(周崇莲),et al.Researches on improving the quality of forest land and the productivity of artificial Cunninghamia lanceolata stands.Chinese Journal of Applied Ecology(应用生态学报),1990,1(2):97-106(in Chinese)
[25]Chen L-C(陈龙池),Wang S-L(汪思龙),Chen C-Y(陈楚莹).Degradation mechanism of Chinese fir plantation.Chinese Journal of Applied Ecology(应用生态学报),2004,15(10):1953-1957(in Chinese)
[26]Fang Q(方奇).Effects of continued plating of Chinese fir on the fertility of soil and the growth of stands.Scientia Silvae Sinicae(林业科学),1987,23(4):389-396(in Chinese)
[27]Ma X-Q(马祥庆),Liu A-Q(刘爱琴),Ma Z(马壮),et al.A comparative study on nutrient accumulation and distribution of different generations of Chinese fir plantations.Chinese Journal of Applied Ecology(应用生态学报),2000,11(4):501-506(in Chinese)
[28]Lin S-Z(林思祖),Huang S-G(黄世国),Cao G-Q(曹光球),et al.Autointoxication of Chinese fir.Chinese Journal of Applied Ecology(应用生态学报),1999,10(6):661-664(in Chinese)
[29]Lin T-L(林同龙).Application effect research on intimate natural forestry management techniques for Cunninghamia lanceolata plantation.Journal of Central South University of Forestry&Technology(中南林业科技大学学报),2012,32(3):11-16(in Chinese)
[30]Lin K-M(林开敏),Huang B-L(黄宝龙).Studies onβ-diversity index of undergrowth plant in Chinese fir plantation.Biodiversity Science(生物多样性),2001,9(2):157-161(in Chinese)
[31]Lin K-M(林开敏),Yu X-T(俞新妥),Huang B-L(黄宝龙),et al.Dynamical characteristics of undergrowth plant diversity in Chinese fir plantation.Chinese Journal of Applied&Environmental Biology(应用与环境生物学报),2001,7(1):13-19(in Chinese)
[32]Fan S-H(范少辉),Ma X-Q(马祥庆),Fu R-S(傅瑞树),et al.Comparative study on underground vegetation develop of different generation plantations of Chinese fir.Forest Research(林业科学研究),2001,14(1):8-16(in Chinese)
[33]Bailey JD,Covington WW.Evaluating ponderosa pine regeneration rates following ecological restoration treatments in northern Arizona,USA.Forest Ecology and Management,2002,155:271-278
[34]Zhu JJ,Mao ZH,Hu LL,et al.Plant diversity of secondary forests in response to anthropogenic disturbance levels in montane regions of northeastern China.Journal of Forest Research,2007,12:403-416
[35]Pammenter NW,Farrant JM,Berjak P.Recalcitrant seeds:Short-term storage effects in Avicennia marina(Forsk.)Vierh.may be germination-associated.Annals of Botany,1984,54:843-846
[36]Gaucher C,Gougeon S,Mauffette Y,et al.Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple(Acer saccharum)and yellow birch(Betula alleghaniensis)seedlings.Tree Physiology,2005,25:93-100
[37]Yan X-F(闫兴富),Wang J-L(王建礼),Zhou L-B(周立彪).Effects of light intensity on Quercus liaotungensis seed germination and seedling growth.Chinese Journal of Applied Ecology(应用生态学报),2011,22(7):1682-1688(in Chinese)
[38]Yan X-F(闫兴富),Zhou L-B(周立彪),Du Q(杜茜).Effects of different shading treatments on the germination of Fraxinus chinensis seed.Journal of Northwest Forestry University(西北林学院学报),2009,24(4):89-92(in Chinese)
[39]Xu N(许宁),Han H-Y(憨宏艳),Gan X-H(甘小洪).Effects of light and ground cover on seed germination and seedling initial growth of Tetracentron sinense.Journal of Plant Resources and Environment(植物资源与环境学报),2015,24(3):85-93(in Chinese)
[40]Xu Z-Q(许中旗),Huang X-R(黄选瑞),Xu C-L(徐成立),et al.The impacts of light conditions on the growth and morphology of Quercus mongolica seedlings.Acta Ecologica Sinica(生态学报),2009,29(3):1121-1128(in Chinese)
[41]Mokany K,Raison R,Prokushkin AS.Critical analysis of root:shoot ratios in terrestrial biomes.Global Change Biology,2006,12:84-96
[42]Yan X-F(闫兴富),Cao M(曹敏).Effects of forest gap on the seedling growth of a canopy tree species Pometia tomentosa in tropical rainforest of Xishuangbanna.Chinese Journal of Applied Ecology(应用生态学报),2008,19(2):238-244(in Chinese)
[43]Hong M(洪明),Guo Q-S(郭泉水),Nie B-H(聂必红),et al.Responses of Cynodon dactylon population in hydro-fluctuation belt of Three Gorges Reservoir area to flooding-drying habitat change.Chinese Journal of Applied Ecology(应用生态学报),2011,22(11):2829-2835(in Chinese)
[44]Ma Z-L(马志良),Yang W-Q(杨万勤),Wu F-Z(吴福忠),et al.Effects of shading on the aboveground biomass and stiochiometry characteristics of Medicago sativa.Chinese Journal of Applied Ecology(应用生态学报),2014,25(11):3139-3144(in Chinese)
[45]Verdu M,Garcia Fayos P.Nucleation processes in a Mediterranean bird-dispersed plant.Functional Ecology,1996,10:275-280
[46]Xu F(徐飞),Guo W-H(郭卫华),Xu W-H(徐伟红),et al.Effects of light intensity on growth and photosynthesis of seedlings of Quercus acutissima and Robinia pseudoacacia.Acta Ecologica Sinica(生态学报),2010,30(12):3098-3107(in Chinese)
[47]Mediavilla S,Escudero A.Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water.European Journal of Forest Research,2010,129:697-706
[48]Valladares F,Niinemets.Shade tolerance,a key plant feature of complex nature and consequences.Annual Review of Ecology,Evolution,and Systematics,2008,39:237-257
[49]Wright IJ,Reich PB,Westoby M,et al.The worldwide leaf economics spectrum.Nature,2004,428:821-827
[50]Walters MB,Reich PB.Are shade tolerance,survival,and growth linked?Low light and nitrogen effects on hardwood seedlings.Ecology,1996,77:841-853
[2]Du XJ,Guo QF,Gao XM,et al.Seed rain,soil seed bank,seed loss and regeneration of Castanopsis fargesii(Fagaceae)in a subtropical evergreen broad-leaved forest.Forest Ecology and Management,2007,238:212-219
[3]Scholes J,Press M,Zipperlen S.Differences in light energy utilisation and dissipation between dipterocarp rain forest tree seedlings.Oecologia,1996,109:41-48
[4]Han Y-Z(韩有志),Wang Z-Q(王政权).Spatial heterogeneity and forest regeneration.Chinese Journal of Applied Ecology(应用生态学报),2002,13(5):615-619(in Chinese)
[5]Nicotra AB,Chazdon RL,Iriarte SVB.Spatial heterogeneity of light and woody seedling regeneration in tropical wet forests.Ecology,1999,80:1908-1926
[6]Diaci J.Regeneration dynamics in a Norway spruce plantation on a silver fir-beech forest site in the Slovenian Alps.Forest Ecology and Management,2002,161:27-38
[7]Lai J-S(赖江山),Li Q-M(李庆梅),Xie Z-Q(谢宗强).Seed germinating characteristics of the endangered plant Abies chensiensis.Chinese Journal of Plant Ecology(植物生态学报),2003,27(5):661-666(in Chinese)
[8]Wu Y(吴彦),Liu Q(刘庆),He H(何海),et al.Effects of light and temperature on seed germination of Picea asperata and Betula albo-sinensis.Chinese Journal of Applied Ecology(应用生态学报),2004,15(12):2229-2232(in Chinese)
[9]Yan X-F(闫兴富),Cao M(曹敏).Effects of light intensity on seed germination and seedling early growth of Shorea wantianshuea.Chinese Journal of Applied Ecology(应用生态学报),2007,18(1):23-29(in Chinese)
[10]Benigno GR,Mulualem T,Guillermo,et al.Seed germination and seedling establishment of Neotropical dry forest species in response to temperature and light conditions.Journal of Forestry Research,2009,20:99-104
[11]Mc Kinnon LM,Mitchell AK.Photoprotection,not increased growth,characterizes the response of Engelmann spruce(Picea engelmannii)seedlings to high light,even when resources are plentiful.New Phytologist,2003,160:69-79
[12]Mc Connaughay KDM,Coleman JS.Biomass allocation in plants:Ontogeny or optimality?A test along three resource gradients.Ecology,1999,80:2581-2593
[13]Grubb PJ,Lee WG,Kollmann J,et al.Interaction of irradiance and soil nutrient supply on growth of seedlings of ten European tall-shrub species and Fagus sylvatica.Journal of Ecology,1996,84:827-840
[14]Evans JR,Poorter H.Photosynthetic acclimation of plants to growth irradiance:The relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain.Plant,Cell&Environment,2001,24:755-767
[15]Mokany K,Raison R,Prokushkin AS.Critical analysis of root:Shoot ratios in terrestrial biomes.Global Change Biology,2006,12:84-96
[16]Lockhart BR,Gardiner ES,Hodges JD,et al.Carbon allocation and morphology of cherrybark oak seedlings and sprouts under three light regimes.Annals of Forest Science,2008,65:801
[17]Stearns SC.The Evolution of Life Histories.Oxford:Oxford University Press,1992
[18]Bloom AJ,Chapin FS,Mooney HA.Resource limitation in plants:An economic analogy.Annual Review of Ecology and Systematics,1985,16:363-392
[19]Geng H-L(耿浩林),Wang Y-H(王玉辉),Wang FY(王风玉),et al.The dynamics of root-shoot ratio and its environmental effective factors of recovering Leymus chinensis steppe vegetation in Inner Mongolia.Acta Ecologica Sinica(生态学报),2008,28(10):4629-4634(in Chinese)
[20]Zhang M(张敏).The Germination,Seedling Survival and Early Growth of the Main Tree Species in Response to Light Regimes in Secondary Forests,Northeast China.Ph D Thesis.Shenyang:Institute Appled Ecology,Chinese Academy of Sciences,2012(in Chinese)
[21]Portsmuth A,Niinemets.Structural and physiological plasticity in response to light and nutrients in five temperate deciduous woody species of contrasting shade tolerance.Functional Ecology,2007,21:61-77
[22]Yu X-T(俞新妥).A summary of the studies on Chinese fir in 2000-2005.Ⅰ.The research development on physiological ecology of Chinese fir.Journal of Fujian College of Forestry(福建林学院学报),2006,26(2):177-185(in Chinese)
[23]Yu X-T(俞新妥).Summary of the studies on the plantation productivity and nutrient cycling in Chinese fir plantation ecosystem.World Forestry Research(世界林业研究),1993,2(3):80-86(in Chinese)
[24]Chen C-Y(陈楚莹),Zhang J-W(张家武),Zhou C-L(周崇莲),et al.Researches on improving the quality of forest land and the productivity of artificial Cunninghamia lanceolata stands.Chinese Journal of Applied Ecology(应用生态学报),1990,1(2):97-106(in Chinese)
[25]Chen L-C(陈龙池),Wang S-L(汪思龙),Chen C-Y(陈楚莹).Degradation mechanism of Chinese fir plantation.Chinese Journal of Applied Ecology(应用生态学报),2004,15(10):1953-1957(in Chinese)
[26]Fang Q(方奇).Effects of continued plating of Chinese fir on the fertility of soil and the growth of stands.Scientia Silvae Sinicae(林业科学),1987,23(4):389-396(in Chinese)
[27]Ma X-Q(马祥庆),Liu A-Q(刘爱琴),Ma Z(马壮),et al.A comparative study on nutrient accumulation and distribution of different generations of Chinese fir plantations.Chinese Journal of Applied Ecology(应用生态学报),2000,11(4):501-506(in Chinese)
[28]Lin S-Z(林思祖),Huang S-G(黄世国),Cao G-Q(曹光球),et al.Autointoxication of Chinese fir.Chinese Journal of Applied Ecology(应用生态学报),1999,10(6):661-664(in Chinese)
[29]Lin T-L(林同龙).Application effect research on intimate natural forestry management techniques for Cunninghamia lanceolata plantation.Journal of Central South University of Forestry&Technology(中南林业科技大学学报),2012,32(3):11-16(in Chinese)
[30]Lin K-M(林开敏),Huang B-L(黄宝龙).Studies onβ-diversity index of undergrowth plant in Chinese fir plantation.Biodiversity Science(生物多样性),2001,9(2):157-161(in Chinese)
[31]Lin K-M(林开敏),Yu X-T(俞新妥),Huang B-L(黄宝龙),et al.Dynamical characteristics of undergrowth plant diversity in Chinese fir plantation.Chinese Journal of Applied&Environmental Biology(应用与环境生物学报),2001,7(1):13-19(in Chinese)
[32]Fan S-H(范少辉),Ma X-Q(马祥庆),Fu R-S(傅瑞树),et al.Comparative study on underground vegetation develop of different generation plantations of Chinese fir.Forest Research(林业科学研究),2001,14(1):8-16(in Chinese)
[33]Bailey JD,Covington WW.Evaluating ponderosa pine regeneration rates following ecological restoration treatments in northern Arizona,USA.Forest Ecology and Management,2002,155:271-278
[34]Zhu JJ,Mao ZH,Hu LL,et al.Plant diversity of secondary forests in response to anthropogenic disturbance levels in montane regions of northeastern China.Journal of Forest Research,2007,12:403-416
[35]Pammenter NW,Farrant JM,Berjak P.Recalcitrant seeds:Short-term storage effects in Avicennia marina(Forsk.)Vierh.may be germination-associated.Annals of Botany,1984,54:843-846
[36]Gaucher C,Gougeon S,Mauffette Y,et al.Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple(Acer saccharum)and yellow birch(Betula alleghaniensis)seedlings.Tree Physiology,2005,25:93-100
[37]Yan X-F(闫兴富),Wang J-L(王建礼),Zhou L-B(周立彪).Effects of light intensity on Quercus liaotungensis seed germination and seedling growth.Chinese Journal of Applied Ecology(应用生态学报),2011,22(7):1682-1688(in Chinese)
[38]Yan X-F(闫兴富),Zhou L-B(周立彪),Du Q(杜茜).Effects of different shading treatments on the germination of Fraxinus chinensis seed.Journal of Northwest Forestry University(西北林学院学报),2009,24(4):89-92(in Chinese)
[39]Xu N(许宁),Han H-Y(憨宏艳),Gan X-H(甘小洪).Effects of light and ground cover on seed germination and seedling initial growth of Tetracentron sinense.Journal of Plant Resources and Environment(植物资源与环境学报),2015,24(3):85-93(in Chinese)
[40]Xu Z-Q(许中旗),Huang X-R(黄选瑞),Xu C-L(徐成立),et al.The impacts of light conditions on the growth and morphology of Quercus mongolica seedlings.Acta Ecologica Sinica(生态学报),2009,29(3):1121-1128(in Chinese)
[41]Mokany K,Raison R,Prokushkin AS.Critical analysis of root:shoot ratios in terrestrial biomes.Global Change Biology,2006,12:84-96
[42]Yan X-F(闫兴富),Cao M(曹敏).Effects of forest gap on the seedling growth of a canopy tree species Pometia tomentosa in tropical rainforest of Xishuangbanna.Chinese Journal of Applied Ecology(应用生态学报),2008,19(2):238-244(in Chinese)
[43]Hong M(洪明),Guo Q-S(郭泉水),Nie B-H(聂必红),et al.Responses of Cynodon dactylon population in hydro-fluctuation belt of Three Gorges Reservoir area to flooding-drying habitat change.Chinese Journal of Applied Ecology(应用生态学报),2011,22(11):2829-2835(in Chinese)
[44]Ma Z-L(马志良),Yang W-Q(杨万勤),Wu F-Z(吴福忠),et al.Effects of shading on the aboveground biomass and stiochiometry characteristics of Medicago sativa.Chinese Journal of Applied Ecology(应用生态学报),2014,25(11):3139-3144(in Chinese)
[45]Verdu M,Garcia Fayos P.Nucleation processes in a Mediterranean bird-dispersed plant.Functional Ecology,1996,10:275-280
[46]Xu F(徐飞),Guo W-H(郭卫华),Xu W-H(徐伟红),et al.Effects of light intensity on growth and photosynthesis of seedlings of Quercus acutissima and Robinia pseudoacacia.Acta Ecologica Sinica(生态学报),2010,30(12):3098-3107(in Chinese)
[47]Mediavilla S,Escudero A.Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water.European Journal of Forest Research,2010,129:697-706
[48]Valladares F,Niinemets.Shade tolerance,a key plant feature of complex nature and consequences.Annual Review of Ecology,Evolution,and Systematics,2008,39:237-257
[49]Wright IJ,Reich PB,Westoby M,et al.The worldwide leaf economics spectrum.Nature,2004,428:821-827
[50]Walters MB,Reich PB.Are shade tolerance,survival,and growth linked?Low light and nitrogen effects on hardwood seedlings.Ecology,1996,77:841-853