红松雌球果发育对枝叶营养生长和生物量积累的影响
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摘要
通过对红松生殖枝和营养枝当年的枝叶生长及生物量积累差异的分析,研究2年生雌球果发育对枝叶营养生长和生物量积累的影响。结果表明:2年生雌球果抑制了1年生雌球果的发育,生殖枝当年的营养生长比营养枝当年的营养生长旺盛;生殖枝当年的生长量(顶枝芽除外)与母枝的枝长和基径呈显著正相关(P<0.05),生殖枝当年的顶枝和侧枝生物量与母枝的枝生物量呈显著正相关(P<0.05); 2年生雌球果数量与生殖枝当年的顶枝基径、侧枝长、侧枝基径、侧枝芽数量和侧枝数量呈显著正相关(P<0.05),2年生雌球果生物量与当年的顶枝和侧枝生物量及总生物量呈显著正相关(P<0.05),说明2年生雌球果的发育促进了生殖枝当年的营养生长和生物量积累;生殖枝当年的总生物量中分配给顶枝、侧枝、顶枝针叶、侧枝针叶的生物量比营养枝多,生殖枝中超过87%的当年生物量分配给2年生雌球果,仅有0.22%的比例分配给1年生雌球果;通径分析表明,2年生雌球果的生物量积累直接影响生殖枝母枝的生物量积累,对生殖枝和营养枝当年生物量积累没有直接影响,而生殖枝母枝的生物量直接影响生殖枝和营养枝当年的生物量积累。
To clarify the effects of 2-year-old female cone's development on vegetative growth and biomass accumulation of shoots and leaves in Pinus koraiensis,the differences of shoot growth and biomass accumulation in the current year were analyzed with reproductive branches and vegetative branches. The results showed that the 2-year-old female cones inhibited the development of1-year-old female cones. The vegetative growth of the reproductive branches in the current year was more vigorous than that of the vegetative branches. The vegetative growth of the reproductive branches( excluding the terminal buds) in the current year was positively correlated with the length and basal diameter of the parent shoots( P<0.05). The biomass of the terminal and lateral shoots of the reproductive branches in the current year was positively related with the biomass of the parent shoots( P<0.05). The number of 2-year-old female cones was significantly positively correlated with the basal diameter of terminal shoot,length of lateral shoot,diameter of lateral shoot,number of lateral shoot buds,and number of lateral shoot of the reproductive branches in the current year( P <0.05). The biomass of 2-year-old female cones was significantly positively correlated with the biomass of terminal shoots,the biomass of lateral shoots,and the biomass of total shoots in the current year( P<0.05). These results indicated that the development of 2-yearold female cones could promote vegetative growth and biomass accumulation in the current year's growth of the reproductive branches. The terminal shoots and needles,the lateral shoots and needles of the reproductive branches were allocated more biomass in the current year than the vegetative branch. More than 87% of the biomass in the reproductive branches was allocated to 2-year-old female cones,while only 0.22% was allocated to 1-year-old female cones. The development of 2-year-old female cones directly affected the biomass accumulation of the parent shoots of the reproductive branches,while the biomass of the parent shoots of the reproductive branches directly affected the biomass accumulation of the reproductive branches and vegetative branches in the current year.
To clarify the effects of 2-year-old female cone's development on vegetative growth and biomass accumulation of shoots and leaves in Pinus koraiensis,the differences of shoot growth and biomass accumulation in the current year were analyzed with reproductive branches and vegetative branches. The results showed that the 2-year-old female cones inhibited the development of1-year-old female cones. The vegetative growth of the reproductive branches in the current year was more vigorous than that of the vegetative branches. The vegetative growth of the reproductive branches( excluding the terminal buds) in the current year was positively correlated with the length and basal diameter of the parent shoots( P<0.05). The biomass of the terminal and lateral shoots of the reproductive branches in the current year was positively related with the biomass of the parent shoots( P<0.05). The number of 2-year-old female cones was significantly positively correlated with the basal diameter of terminal shoot,length of lateral shoot,diameter of lateral shoot,number of lateral shoot buds,and number of lateral shoot of the reproductive branches in the current year( P <0.05). The biomass of 2-year-old female cones was significantly positively correlated with the biomass of terminal shoots,the biomass of lateral shoots,and the biomass of total shoots in the current year( P<0.05). These results indicated that the development of 2-yearold female cones could promote vegetative growth and biomass accumulation in the current year's growth of the reproductive branches. The terminal shoots and needles,the lateral shoots and needles of the reproductive branches were allocated more biomass in the current year than the vegetative branch. More than 87% of the biomass in the reproductive branches was allocated to 2-year-old female cones,while only 0.22% was allocated to 1-year-old female cones. The development of 2-year-old female cones directly affected the biomass accumulation of the parent shoots of the reproductive branches,while the biomass of the parent shoots of the reproductive branches directly affected the biomass accumulation of the reproductive branches and vegetative branches in the current year.
引文
钱杨,孙洪刚,董汝湘,等.2018.针叶树碳水化合物分配研究进展.林业科学,54(1):141-153.
沈海龙,丛健,张鹏,等.2011.开敞度调控对次生林林冠下红松径高生长量和地上生物量的影响.应用生态学报,22(11):2781-2791.
王娟,张春雨,赵秀海,等.2011.雌雄异株植物鼠李的生殖分配.生态学报,31(21):6371-6377.
谢宗强,陈伟烈,胡东,等.1998.濒危植物银杉的结实特性及动物对果实的危害性.植物生态学报,22(4):319-326.
殷东生,沈海龙.2016.林木大年结实特征及其影响因素研究进展.生态学杂志,35(2):542-550.
尹红,王靖,刘洪滨,等.2011.小兴安岭红松径向生长对未来气候变化的响应.生态学报,31(24):7343-7350.
张鹏,刘畅,沈海龙.2017.世界食用松资源及其开发利用现状.世界林业研究,30(1):12-17.
张华新.2000.油松雌球果发育对枝条生长的影响.林业科学研究,13(5):505-511.
郑杨,董利虎,李凤日.2016.黑龙江省红松人工林枝条分布数量模拟.应用生态学报,27(7):2172-2180.
Alla AQ,Camarero JJ,Gabriel Montserrat-MartíMM.2012.Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species.Trees,26:841-850.
Brienen RJW,Zuidema PA,During HJ.2006.Autocorrelated growth of tropical forest trees:Unraveling patterns and quantifying consequences.Forest Ecology and Management,237:179-190.
Castillollanque F,Rapoport HF.2011.Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees(Olea europaea L.).Trees,25:823-832.
Chen H,Niklas KJ,Yang DM,et al.2009.The effect of twig architecture and seed number on seed size variation in subtropical woody species.New Phytologist,183:1212-1221.
Crone EE,McIntire EJB,Brodie J.2011.What defines mastseeding?Spatiotemporal patterns of cone production by whitebark pine.Journal of Ecology,99:438-444.
Dick JM,Leakey RR,Jarvis PG.1990.Influence of female cones on the vegetative growth of Pinus contorta trees.Tree Physiology,6:151-163.
Elloumi O,Ghrab M,Mimoun MB.2014.Effects of flower buds removal on seasonal starch storage and mobilization in fruiting and non-fruiting branches of pistachio trees cv.Mateur under dry and warm climate.Scientia Horticulturae,172:19-25.
Fox JC,Ades PK,Bi H.2001.Stochastic structure and individual-tree growth models.Forest Ecology and Management,154:261-276.
Han Q,Kabeya D,Hoch G.2011.Leaf traits,shoot growth and seed production in mature Fagus sylvatica trees after 8years of CO2enrichment.Annals of Botany,107:1405-1411.
Han Q,Kabeya D.2017.Recent developments in understanding mast seeding in relation to dynamics of carbon and nitrogen resources in temperate trees.Ecological Research,32:771-778.
Hoch G.2005.Fruit-bearing branchlets are carbon autonomous in mature broad-leaved temperate forest trees.Plant,Cell and Environment,28:651-659.
Ichie T,Kenzo T,Kitahashi Y,et al.2005.How does Dryobalanops aromatica supply carbohydrate resources for reproduction in a masting year?Trees,19:703-710.
Ishihara MI,Kikuzawa K.2009.Annual and spatial variation in shoot demography associated with masting in Betula grossa:Comparison between mature trees and saplings.Annals of Botany,104:1195-1205.
Martín D,Vazquez-PiquéJ,Carevic FS,et al.2015.Trade-off between stem growth and acorn production in holm oak.Trees,29:825-834.
Miyazaki Y,Hiura T,Funada R.2007.Allocation of photo-assimilated13C from reproductive and non-reproductive shoots to fruits in Styrax obassia.Plant Species Biology,22:53-57.
Miyazaki Y.2013.Dynamics of internal carbon resources during masting behavior in trees.Ecological Research,28:143-150.
Ne’eman G,Goubitz S,Werger MJA.2011.Relationships between tree size,crown shape,gender segregation and sex allocation in Pinus halepensis,a Mediterranean pine tree.Annals of Botany,108:197-206.
Newell EA.1991.Direct and delayed costs of reproduction in Aesculus californica.Journal of Ecology,79:365-378.
Obeso JR.1997.Costs of reproduction in Ilex aquifolium:Effects at tree branch and leaf levels.Journal of Ecology,85:159-166.
Oitate H,Noguchi K,Sone K,et al.2011.Patterns of photoassimilate translocation to reproductive shoots from adjacent shoots in Camellia sasanqua by manipulation of sink-source balance between the shoots.Journal of Plant Research,124:131-136.
Pearse IS,Koenig WD,Kelly D.2016.Mechanisms of mast seeding:Resources,weather,cues,and selection.New Phytologist,212:546-562.
Pesendorfer MB,Koenig WD,Pearse IS,et al.2016.Individual resource limitation combined with population-wide pollen availability drives masting in the valley oak(Quercus lobata).Journal of Ecology,104:637-645.
Rapp JM,McIntire EJB,Crone EE.2013.Sex allocation,pollen limitation and masting in whitebark pine.Journal of Ecology,101:1345-1352.
Reekie EG,Bazzaz FA.1987.Reproductive effort in plants.II.Does carbon reflect the allocation of other resources?American Naturalist,129:897-906.
Sala A,Hopping K,McIntire EJB,et al.2012.Masting in whitebark pine(Pinus albicaulis)depletes stored nutrients.New Phytologist,196:189-199.
Sanchez-Humanes B,Sork VL,Espelta JM.2011.Trade-offs between vegetative growth and acorn production in Quercus lobata during a mast year:The relevance of crop size and hierarchical level within the canopy.Oecologia,166:101-110.
Sgro CM,Hoffmann AA.2004.Genetic correlations tradeoffs and environmental variation.Heredity,93:241-248.
Yasumura Y.2006.Resource allocation to vegetative and reproductive growth in relation to mast seeding in Fagus crenata.Forest Ecology and Management,229:228-233.
Zhang KM,Li Z,Li Y,et al.2013.Carbohydrate accumulation may be the proximate trigger of anthocyanin biosynthesis under autumn conditions in Begonia semperflorens.Plant Biology,15:991-1000.
Zywiec M,Zielonka T.2013.Does a heavy fruit crop reduce the tree ring increment?Results from a 12-year study in a subalpine zone.Trees,27:1365-1373.
沈海龙,丛健,张鹏,等.2011.开敞度调控对次生林林冠下红松径高生长量和地上生物量的影响.应用生态学报,22(11):2781-2791.
王娟,张春雨,赵秀海,等.2011.雌雄异株植物鼠李的生殖分配.生态学报,31(21):6371-6377.
谢宗强,陈伟烈,胡东,等.1998.濒危植物银杉的结实特性及动物对果实的危害性.植物生态学报,22(4):319-326.
殷东生,沈海龙.2016.林木大年结实特征及其影响因素研究进展.生态学杂志,35(2):542-550.
尹红,王靖,刘洪滨,等.2011.小兴安岭红松径向生长对未来气候变化的响应.生态学报,31(24):7343-7350.
张鹏,刘畅,沈海龙.2017.世界食用松资源及其开发利用现状.世界林业研究,30(1):12-17.
张华新.2000.油松雌球果发育对枝条生长的影响.林业科学研究,13(5):505-511.
郑杨,董利虎,李凤日.2016.黑龙江省红松人工林枝条分布数量模拟.应用生态学报,27(7):2172-2180.
Alla AQ,Camarero JJ,Gabriel Montserrat-MartíMM.2012.Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species.Trees,26:841-850.
Brienen RJW,Zuidema PA,During HJ.2006.Autocorrelated growth of tropical forest trees:Unraveling patterns and quantifying consequences.Forest Ecology and Management,237:179-190.
Castillollanque F,Rapoport HF.2011.Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees(Olea europaea L.).Trees,25:823-832.
Chen H,Niklas KJ,Yang DM,et al.2009.The effect of twig architecture and seed number on seed size variation in subtropical woody species.New Phytologist,183:1212-1221.
Crone EE,McIntire EJB,Brodie J.2011.What defines mastseeding?Spatiotemporal patterns of cone production by whitebark pine.Journal of Ecology,99:438-444.
Dick JM,Leakey RR,Jarvis PG.1990.Influence of female cones on the vegetative growth of Pinus contorta trees.Tree Physiology,6:151-163.
Elloumi O,Ghrab M,Mimoun MB.2014.Effects of flower buds removal on seasonal starch storage and mobilization in fruiting and non-fruiting branches of pistachio trees cv.Mateur under dry and warm climate.Scientia Horticulturae,172:19-25.
Fox JC,Ades PK,Bi H.2001.Stochastic structure and individual-tree growth models.Forest Ecology and Management,154:261-276.
Han Q,Kabeya D,Hoch G.2011.Leaf traits,shoot growth and seed production in mature Fagus sylvatica trees after 8years of CO2enrichment.Annals of Botany,107:1405-1411.
Han Q,Kabeya D.2017.Recent developments in understanding mast seeding in relation to dynamics of carbon and nitrogen resources in temperate trees.Ecological Research,32:771-778.
Hoch G.2005.Fruit-bearing branchlets are carbon autonomous in mature broad-leaved temperate forest trees.Plant,Cell and Environment,28:651-659.
Ichie T,Kenzo T,Kitahashi Y,et al.2005.How does Dryobalanops aromatica supply carbohydrate resources for reproduction in a masting year?Trees,19:703-710.
Ishihara MI,Kikuzawa K.2009.Annual and spatial variation in shoot demography associated with masting in Betula grossa:Comparison between mature trees and saplings.Annals of Botany,104:1195-1205.
Martín D,Vazquez-PiquéJ,Carevic FS,et al.2015.Trade-off between stem growth and acorn production in holm oak.Trees,29:825-834.
Miyazaki Y,Hiura T,Funada R.2007.Allocation of photo-assimilated13C from reproductive and non-reproductive shoots to fruits in Styrax obassia.Plant Species Biology,22:53-57.
Miyazaki Y.2013.Dynamics of internal carbon resources during masting behavior in trees.Ecological Research,28:143-150.
Ne’eman G,Goubitz S,Werger MJA.2011.Relationships between tree size,crown shape,gender segregation and sex allocation in Pinus halepensis,a Mediterranean pine tree.Annals of Botany,108:197-206.
Newell EA.1991.Direct and delayed costs of reproduction in Aesculus californica.Journal of Ecology,79:365-378.
Obeso JR.1997.Costs of reproduction in Ilex aquifolium:Effects at tree branch and leaf levels.Journal of Ecology,85:159-166.
Oitate H,Noguchi K,Sone K,et al.2011.Patterns of photoassimilate translocation to reproductive shoots from adjacent shoots in Camellia sasanqua by manipulation of sink-source balance between the shoots.Journal of Plant Research,124:131-136.
Pearse IS,Koenig WD,Kelly D.2016.Mechanisms of mast seeding:Resources,weather,cues,and selection.New Phytologist,212:546-562.
Pesendorfer MB,Koenig WD,Pearse IS,et al.2016.Individual resource limitation combined with population-wide pollen availability drives masting in the valley oak(Quercus lobata).Journal of Ecology,104:637-645.
Rapp JM,McIntire EJB,Crone EE.2013.Sex allocation,pollen limitation and masting in whitebark pine.Journal of Ecology,101:1345-1352.
Reekie EG,Bazzaz FA.1987.Reproductive effort in plants.II.Does carbon reflect the allocation of other resources?American Naturalist,129:897-906.
Sala A,Hopping K,McIntire EJB,et al.2012.Masting in whitebark pine(Pinus albicaulis)depletes stored nutrients.New Phytologist,196:189-199.
Sanchez-Humanes B,Sork VL,Espelta JM.2011.Trade-offs between vegetative growth and acorn production in Quercus lobata during a mast year:The relevance of crop size and hierarchical level within the canopy.Oecologia,166:101-110.
Sgro CM,Hoffmann AA.2004.Genetic correlations tradeoffs and environmental variation.Heredity,93:241-248.
Yasumura Y.2006.Resource allocation to vegetative and reproductive growth in relation to mast seeding in Fagus crenata.Forest Ecology and Management,229:228-233.
Zhang KM,Li Z,Li Y,et al.2013.Carbohydrate accumulation may be the proximate trigger of anthocyanin biosynthesis under autumn conditions in Begonia semperflorens.Plant Biology,15:991-1000.
Zywiec M,Zielonka T.2013.Does a heavy fruit crop reduce the tree ring increment?Results from a 12-year study in a subalpine zone.Trees,27:1365-1373.