垂直切根对当年羊草构件及繁殖资源分配的影响
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摘要
为探究羊草(Leymus chinensis)在外界环境干扰下的繁殖分配,本研究在羊草不同发育时期(返青期、孕穗期、抽穗初期)进行不同间距(10、20、30cm)的切根处理,对羊草叶、茎、穗、根茎的生物量以及营养元素含量(全氮、全磷)、能量累积和繁殖分配进行比较分析。结果表明,切根对羊草穗的生物量、全氮、全磷、能量等资源的繁殖分配影响显著(P<0.05),同一时期切根,羊草穗的繁殖分配表现为切根间距30cm>20cm>10cm;同一切根间距,羊草穗的繁殖分配为:抽穗初期>孕穗期>返青期;切根后羊草叶、茎、穗的全氮含量显著升高(P<0.05)。返青期切根间距10cm处理羊草根茎的资源分配比例增加,促进了羊草的无性繁殖,而抽穗初期切根间距30cm羊草穗的繁殖分配比例增加,是利于羊草有性繁殖的最佳切根处理。综上所述,垂直切根可以影响羊草繁殖分配,羊草的繁殖分配是其应对环境干扰的重要调节机制。
To investigate the reproductive allocation of Leymus chinensis under the interference of the external environment,root cutting treatments with 10,20,and 30 cm spacing were conducted at different growth periods(regreening,booting,early heading).The biomass of leaves,stems,panicles,and rhizomes and total N and P content were measured.Results showed that root cutting significantly influenced the reproductive distribution of panicle biomass,total N,and total P.Within the same growth period,panicle reproductive distribution was highest in the 30 cm treatment and lowest in the 10 cm treatment.Within the same root cutting treatment,reproductive allocation of L.chinensis was highest for early heading and lowest for regreening growth periods.Total N content of L.chinensis leaves,stems,and panicles was significantly higher after root cutting(P<0.05).In the 10 cm root cutting treatment,the proportion of rhizomes increased because of the initiation of asexual reproduction.The proportion of panicles increased under the 30 cm root cutting treatment,indicating that this was the optimal spacing for sexual reproduction of L.chinensis.Results of this study show that vertical root cutting affects the reproductive distribution of L.chinensis and is therefore an important regulation mechanism in response to environmental disturbance.
To investigate the reproductive allocation of Leymus chinensis under the interference of the external environment,root cutting treatments with 10,20,and 30 cm spacing were conducted at different growth periods(regreening,booting,early heading).The biomass of leaves,stems,panicles,and rhizomes and total N and P content were measured.Results showed that root cutting significantly influenced the reproductive distribution of panicle biomass,total N,and total P.Within the same growth period,panicle reproductive distribution was highest in the 30 cm treatment and lowest in the 10 cm treatment.Within the same root cutting treatment,reproductive allocation of L.chinensis was highest for early heading and lowest for regreening growth periods.Total N content of L.chinensis leaves,stems,and panicles was significantly higher after root cutting(P<0.05).In the 10 cm root cutting treatment,the proportion of rhizomes increased because of the initiation of asexual reproduction.The proportion of panicles increased under the 30 cm root cutting treatment,indicating that this was the optimal spacing for sexual reproduction of L.chinensis.Results of this study show that vertical root cutting affects the reproductive distribution of L.chinensis and is therefore an important regulation mechanism in response to environmental disturbance.
引文
[1]王殿才,郑金艳,李凤兰.松土补播改良退化草场试验研究.畜牧兽医科技信息,2009(12):277-286.WANG D C,ZHENG J Y,LI F L.Experimental study on the improvement of degraded grassland by sowing and sowing.Chinese Journal of Animal Husbandry and Veterinary Medicine,2009(12):277-286.
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[4]张俊刚,姜慧敏,李雅茹,宝音陶格涛.施肥和灌溉对切根改良草地群落的影响.中国草地学报,2016,38(1):81-86.ZHANG J G,JIANG H M,LI Y R,Baoyintaogetao.Effect of fertilizing and irrigation on grassland improved by cutting root.Chinese Journal of Grassland,2016,38(1):81-86.
[5]宝音陶格涛.不同改良措施下退化羊草草原群落恢复演替阶段规律研究.呼和浩特:内蒙古大学博士学位论文,2009.Baoyintaogetao.The study on dynamice succession of community in degraded steppe of Leymus chinensis after different omprovement measures.PhD Thesis.Hohhot:Inner Mongolia University,2009.
[6]张佳良,曹杰,邓波,陈积山,张月学.垂直切根可以改善松嫩草地羊草生长.草业科学,2017,34(5):1057-1063.ZHANG J L,CAO J,DENG B,CHEN J S,ZHANG Y X.Root cutting can improve the growth of Leymus chinensis in Songnen Plain.Pratacultural Science,2017,34(5):1057-1063.
[7]乌仁其其格,闫瑞瑞,辛晓平,斯钦高娃.切根改良对退化草地羊草群落的影响.内蒙古农业大学学报,2011,32(4):55-58.Wurenqiqige,YAN R R,XIN X P,Siqingaowa.Influence of root cutting on degraded grassland Leymus chiensis community.Journal of Inner Mongolia Agricultural University,2011,32(4):55-58.
[8]袁洪方,王德成,王光辉,刘鹏军,张新田,王明亮.山地草甸类草场破土切根试验研究//2010国际农业工程大会论文集.上海:中国农业机械学会,2010:7249-7253.YUAN H F,WANG D C,WANG G H,LIU P J,ZHANG X T,WANG M L.Research experiment of soil-gashing and root-cutting on mountain meadow steppe soil//2010Proceedings of the International Conference on Agricultural Engineering.Shanghai:China Agricultural Machinery Society,2010:7249-7253.
[9]高志成,田佳妮,霍艳双,舒锴,宝音陶格涛.切根和浅耕翻措施对退化草地生长季土壤性质及植物群落的影响.生态学报,2017,37(11):1-6.GAO Z C,TIAN J N,HUO Y S,SHU K,Baoyintaogetao.Effects of root pruning and shallow plowing on soil properties and plant commmunities in deteriorated steppe.Acta Ecologica Sinica,2017,37(11):1-6.
[10]刘库,谢应忠.不同改良措施的研究综述.中国草地学报,2005,27(5):53-57.LIU K,XIE Y Z.Review on the research of different restorative measures for degraded vegetation.Gressland of China,2005,27(5):53-57.
[11]尤泳,王光辉,张洋,侯丽华.破土切根对退化羊草草地土壤理化特性的影响//农区草业论坛文集.厦门:中国草学会,2008:509-515.YOU Y,WANG G H,ZHANG Y,HOU L H.Effects of soil-gashing and root-cutting on soil physical and chemical properties in Leymus chinensis grassland//Collections of Agricultural grassland forum.Xiamen:China Grassland Society,2008:509-515.
[12]钟章成.植物种群的繁殖对策.生态学杂志,1995,14(1):37-42.ZHONG Z C.Reproductive strategy of plant population.Chinese Journal of Ecology,1995,14(1):37-42.
[13]周纪纶.植物种群生态学.北京:中国高等教育出版社,1992.ZHOU J L.Plant Population Ecology.Beijing:China Higher Education Press,1992.
[14] ORDEN J.The reproductive strategy of Tussilago farfara L.Journal of Ecology,1974,62:291-324.
[15] SAKAI S.Optimal resource allocation to vegetative and sexual reproduction of a plant growing in a spatially varying environment.Journal of Theoretical Biology,1995,175:271-282.
[16] GARDNER S N,MANGEL M.Modeling investments in seed,clonal offspring,and translocation in a clonal plant.Ecology,1999,80:1202-1220.
[17] ABRAHAMSON W G.Demography and vegetative reproduction.//SOLBRIG O T(ed).Demography and Evolution in Plant Populations.Oxford:Blackwell,1980:89-106.
[18] COOK R E.Growth and development in clonal plant populations.//JACKSON J B C,BUSS L W,COOK R E(eds).Population Biology and Evolution of Clonal Organisms.New Haven:Yale University Press,1985:259-296.
[19] ERIKSSON O.Clonal life histories and the evolution of seed recruitment.//DE KROON H,VAN GROENENDAEL J(eds).The Ecology and Evolution of Clonal Plants.Leiden:Backhuys Publisher,1997:211-226.
[20]张文军.羊草克隆整合对碱环境异质性的响应.北京:中国农业大学博士学位论文,2014:89.ZANG W J.Response of Leymus chinensis clonal integration to heterogeneous alkaline environment.PhD Thesis.Beijing:China Agricultural University,2014:89.
[21] SILVERTON J W.Introduction to Plant Population Biology.London:Blackwell Scientific Publications,1993.
[22]李冰,刘左军,赵志刚,胡春,任红梅,伍国强.海拔对钝裂银莲花不同花色居群间繁殖特征及繁殖分配的影响.草业学报,2013,22(1):10-19.LI B,LIU Z J,ZHAO Z G,HU C,REN H M,WU G Q.Influence of altitude on reproductive traits and reproductive allocation of different colours in Anemone obtusiloba populations.Acta Prataculturae Sinica,2013,22(1):10-19.
[23] CHEN X S,LI Y F,XIE Y H,DENG Z M,LI X,LI F,HOU Z Y.Trade-off between allocation to reproductive ramets and rhizome buds in Carex brevicuspis populations along a small-scale elevational gradient.Scientific Reports,2015,5:12688.
[24] XIAO Y,TANG J B,QING H.Trade-offs among growth,clonal,and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration.Hydrobiologia,2011,658:353-363.
[25] FARNWORTH E,BAZZAZ F.Inter-and intra-generic in growth,reproduction and fitness of nine herbaceous annual species in elevated CO2environments.Oecologia,1995,104(4):454-466.
[26] LIU F,CHEN J M,WANG Q F.Trade-offs between sexual and asexual reproduction in a monoecious species Sagittaria pygmaea(Alismataceae):The effect of different nutrient levels.Plant Systematics&Evolution,2009,277(1/2):61-65.
[27] THOMPSON F L,ECKERT C G.Trade-offs between sexual and clonal reproduction in an aquatic plant:Experimental manipulations vs.phenotypic correlations.Journal of Evolutionary Biology,2004,17(3):581-592.
[28] JUERG S,RITA R,ANDREAS E,PETER S.The relative importance of sexual and clonal reproduction for population growth in the perennial herb Fragaria vesca.Population Ecology,2012,54(3):369-380.
[29]杜利霞,李青丰,董宽虎.放牧强度对短花针茅草原冷蒿繁殖特性的影响.草地学报,2007,15(4):367-370.DU L X,LI Q F,DONG K H.Effects of grazing intensity on reproductive characteristics of Artemisia frigida willd.in Stipa breviflora Griseb.Steppe.Acta Agrestia Sinica,2007,15(4):367-370.
[30] VAN GROENENDAEL J M,HABEKOTTE B.Cyperus esculentus L.biology,population dynamics,and possibilities to control this neophyte.Zeitschrift Für Pflanzenkrankheiten Und Pflanzenschutz,.1988(6):61-69.
[31] LOEHLE C.Partitioning of reproductive effort in clonal plants:A benefit-cost model.Oikos,1987,49(2):199-208.
[32] HERBEN T.Coexistence and pattern diversity in communities of clonal organisms:A model based on cellular automata.Abstracta Botanica,1992,16:49-54.
[33]王仁忠,祖元刚,聂绍荃.羊草种群生物量生殖分配的初步研究.应用生态学报,1999,10(5):553-555.WANG R Z,ZU Y G,NIE S Q.Preliminary studty on biomass reproductive allocation in Leymus chinensis population.Chinese Journal of Applied Ecology,1999,10(5):553-555.
[34]陈玲,乌江雨,杨持.不同群落类型对羊草种群生殖分配的影响.内蒙古大学学报,1996,27(5):713-716.CHEN L,WU J Y,YANG C.The influence of different habttat on the reproductive allocation of Leymus chinensis population.Acta Scientiarum Naturalium Universitatis NeiMonggol,1996,27(5):713-716.
[35]王仁忠,祖元刚.羊草种群生物量和能量生殖分配的研究.植物研究,2001,21(2):299-303.WANG R Z,ZU Y G.Biomass and energy allocation in Leymus chinensis population.Bulletin of Botanical Research,2001,21(2):299-303.
[2] ZHANG W J,YANG G W,SUN J J,CHEN J S,ZHANG.Y J.Clonal integration enhances the performance of a clonal plant species under soil alkalinity stress.PloS ONE,2015,10(3):e119942.
[3]邓庆华,李化丹,于翔,乌仁其其格.切根、施肥对羊草营养成分的影响.黑龙江畜牧兽医,2018(2):142-143.DENG Q H,LI H D,YU X,Wurenqiqige.Effects of cut root and fertilization on nutrients of Leymus chinensis.Heilongjiang Animal Science and Veterinary Medicine,2018(2):142-143.
[4]张俊刚,姜慧敏,李雅茹,宝音陶格涛.施肥和灌溉对切根改良草地群落的影响.中国草地学报,2016,38(1):81-86.ZHANG J G,JIANG H M,LI Y R,Baoyintaogetao.Effect of fertilizing and irrigation on grassland improved by cutting root.Chinese Journal of Grassland,2016,38(1):81-86.
[5]宝音陶格涛.不同改良措施下退化羊草草原群落恢复演替阶段规律研究.呼和浩特:内蒙古大学博士学位论文,2009.Baoyintaogetao.The study on dynamice succession of community in degraded steppe of Leymus chinensis after different omprovement measures.PhD Thesis.Hohhot:Inner Mongolia University,2009.
[6]张佳良,曹杰,邓波,陈积山,张月学.垂直切根可以改善松嫩草地羊草生长.草业科学,2017,34(5):1057-1063.ZHANG J L,CAO J,DENG B,CHEN J S,ZHANG Y X.Root cutting can improve the growth of Leymus chinensis in Songnen Plain.Pratacultural Science,2017,34(5):1057-1063.
[7]乌仁其其格,闫瑞瑞,辛晓平,斯钦高娃.切根改良对退化草地羊草群落的影响.内蒙古农业大学学报,2011,32(4):55-58.Wurenqiqige,YAN R R,XIN X P,Siqingaowa.Influence of root cutting on degraded grassland Leymus chiensis community.Journal of Inner Mongolia Agricultural University,2011,32(4):55-58.
[8]袁洪方,王德成,王光辉,刘鹏军,张新田,王明亮.山地草甸类草场破土切根试验研究//2010国际农业工程大会论文集.上海:中国农业机械学会,2010:7249-7253.YUAN H F,WANG D C,WANG G H,LIU P J,ZHANG X T,WANG M L.Research experiment of soil-gashing and root-cutting on mountain meadow steppe soil//2010Proceedings of the International Conference on Agricultural Engineering.Shanghai:China Agricultural Machinery Society,2010:7249-7253.
[9]高志成,田佳妮,霍艳双,舒锴,宝音陶格涛.切根和浅耕翻措施对退化草地生长季土壤性质及植物群落的影响.生态学报,2017,37(11):1-6.GAO Z C,TIAN J N,HUO Y S,SHU K,Baoyintaogetao.Effects of root pruning and shallow plowing on soil properties and plant commmunities in deteriorated steppe.Acta Ecologica Sinica,2017,37(11):1-6.
[10]刘库,谢应忠.不同改良措施的研究综述.中国草地学报,2005,27(5):53-57.LIU K,XIE Y Z.Review on the research of different restorative measures for degraded vegetation.Gressland of China,2005,27(5):53-57.
[11]尤泳,王光辉,张洋,侯丽华.破土切根对退化羊草草地土壤理化特性的影响//农区草业论坛文集.厦门:中国草学会,2008:509-515.YOU Y,WANG G H,ZHANG Y,HOU L H.Effects of soil-gashing and root-cutting on soil physical and chemical properties in Leymus chinensis grassland//Collections of Agricultural grassland forum.Xiamen:China Grassland Society,2008:509-515.
[12]钟章成.植物种群的繁殖对策.生态学杂志,1995,14(1):37-42.ZHONG Z C.Reproductive strategy of plant population.Chinese Journal of Ecology,1995,14(1):37-42.
[13]周纪纶.植物种群生态学.北京:中国高等教育出版社,1992.ZHOU J L.Plant Population Ecology.Beijing:China Higher Education Press,1992.
[14] ORDEN J.The reproductive strategy of Tussilago farfara L.Journal of Ecology,1974,62:291-324.
[15] SAKAI S.Optimal resource allocation to vegetative and sexual reproduction of a plant growing in a spatially varying environment.Journal of Theoretical Biology,1995,175:271-282.
[16] GARDNER S N,MANGEL M.Modeling investments in seed,clonal offspring,and translocation in a clonal plant.Ecology,1999,80:1202-1220.
[17] ABRAHAMSON W G.Demography and vegetative reproduction.//SOLBRIG O T(ed).Demography and Evolution in Plant Populations.Oxford:Blackwell,1980:89-106.
[18] COOK R E.Growth and development in clonal plant populations.//JACKSON J B C,BUSS L W,COOK R E(eds).Population Biology and Evolution of Clonal Organisms.New Haven:Yale University Press,1985:259-296.
[19] ERIKSSON O.Clonal life histories and the evolution of seed recruitment.//DE KROON H,VAN GROENENDAEL J(eds).The Ecology and Evolution of Clonal Plants.Leiden:Backhuys Publisher,1997:211-226.
[20]张文军.羊草克隆整合对碱环境异质性的响应.北京:中国农业大学博士学位论文,2014:89.ZANG W J.Response of Leymus chinensis clonal integration to heterogeneous alkaline environment.PhD Thesis.Beijing:China Agricultural University,2014:89.
[21] SILVERTON J W.Introduction to Plant Population Biology.London:Blackwell Scientific Publications,1993.
[22]李冰,刘左军,赵志刚,胡春,任红梅,伍国强.海拔对钝裂银莲花不同花色居群间繁殖特征及繁殖分配的影响.草业学报,2013,22(1):10-19.LI B,LIU Z J,ZHAO Z G,HU C,REN H M,WU G Q.Influence of altitude on reproductive traits and reproductive allocation of different colours in Anemone obtusiloba populations.Acta Prataculturae Sinica,2013,22(1):10-19.
[23] CHEN X S,LI Y F,XIE Y H,DENG Z M,LI X,LI F,HOU Z Y.Trade-off between allocation to reproductive ramets and rhizome buds in Carex brevicuspis populations along a small-scale elevational gradient.Scientific Reports,2015,5:12688.
[24] XIAO Y,TANG J B,QING H.Trade-offs among growth,clonal,and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration.Hydrobiologia,2011,658:353-363.
[25] FARNWORTH E,BAZZAZ F.Inter-and intra-generic in growth,reproduction and fitness of nine herbaceous annual species in elevated CO2environments.Oecologia,1995,104(4):454-466.
[26] LIU F,CHEN J M,WANG Q F.Trade-offs between sexual and asexual reproduction in a monoecious species Sagittaria pygmaea(Alismataceae):The effect of different nutrient levels.Plant Systematics&Evolution,2009,277(1/2):61-65.
[27] THOMPSON F L,ECKERT C G.Trade-offs between sexual and clonal reproduction in an aquatic plant:Experimental manipulations vs.phenotypic correlations.Journal of Evolutionary Biology,2004,17(3):581-592.
[28] JUERG S,RITA R,ANDREAS E,PETER S.The relative importance of sexual and clonal reproduction for population growth in the perennial herb Fragaria vesca.Population Ecology,2012,54(3):369-380.
[29]杜利霞,李青丰,董宽虎.放牧强度对短花针茅草原冷蒿繁殖特性的影响.草地学报,2007,15(4):367-370.DU L X,LI Q F,DONG K H.Effects of grazing intensity on reproductive characteristics of Artemisia frigida willd.in Stipa breviflora Griseb.Steppe.Acta Agrestia Sinica,2007,15(4):367-370.
[30] VAN GROENENDAEL J M,HABEKOTTE B.Cyperus esculentus L.biology,population dynamics,and possibilities to control this neophyte.Zeitschrift Für Pflanzenkrankheiten Und Pflanzenschutz,.1988(6):61-69.
[31] LOEHLE C.Partitioning of reproductive effort in clonal plants:A benefit-cost model.Oikos,1987,49(2):199-208.
[32] HERBEN T.Coexistence and pattern diversity in communities of clonal organisms:A model based on cellular automata.Abstracta Botanica,1992,16:49-54.
[33]王仁忠,祖元刚,聂绍荃.羊草种群生物量生殖分配的初步研究.应用生态学报,1999,10(5):553-555.WANG R Z,ZU Y G,NIE S Q.Preliminary studty on biomass reproductive allocation in Leymus chinensis population.Chinese Journal of Applied Ecology,1999,10(5):553-555.
[34]陈玲,乌江雨,杨持.不同群落类型对羊草种群生殖分配的影响.内蒙古大学学报,1996,27(5):713-716.CHEN L,WU J Y,YANG C.The influence of different habttat on the reproductive allocation of Leymus chinensis population.Acta Scientiarum Naturalium Universitatis NeiMonggol,1996,27(5):713-716.
[35]王仁忠,祖元刚.羊草种群生物量和能量生殖分配的研究.植物研究,2001,21(2):299-303.WANG R Z,ZU Y G.Biomass and energy allocation in Leymus chinensis population.Bulletin of Botanical Research,2001,21(2):299-303.