中国沙棘克隆生长对灌水强度的响应规律及其激素调控机制
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摘要
关于中国沙棘克隆生长调节研究目前局限于外在机制,旨在探讨其克隆生长对灌水强度的响应规律及其激素调控的内在机制。结果表明:随着灌水强度的增大,分株生长、克隆繁殖、克隆扩散能力先升后降,IAA(吲哚-3-乙酸)、ZR(玉米素核苷)、GA_3(赤霉酸)含量及其与ABA(脱落酸)的比值先升后降而ABA含量先降后升。同时,分株生长、克隆繁殖、克隆扩散能力与IAA、ZR、GA_3含量以及IAA/ABA、ZR/ABA、GA_3/ABA呈极显著或显著正相关,与ABA含量呈极显著负相关。灌水强度过小或过大,IAA、ZR、GA_3含量以及IAA/ABA、ZR/ABA、GA_3/ABA低而ABA含量高,克隆生长潜力受到抑制,种群以分株小、数量少(分布稀疏)、扩散(水平根延伸和分枝)能力弱为特征,克隆生长格局倾向于"游击型"、种群早衰概率高;灌水强度适宜,IAA、ZR、GA_3含量以及IAA/ABA、ZR/ABA、GA_3/ABA高而ABA含量低,克隆生长潜力得以充分发挥,种群以分株大、数量多(分布密集)、扩散能力强为特征,克隆生长格局倾向于"聚集型"、种群稳定性高。随着灌水强度过小-适宜-过大的连续变化,中国沙棘通过改变激素状况调控克隆生长,从而形成与灌水强度相适应的克隆生长格局连续体"游击型-聚集型-游击型",种群稳定性呈"低-高-低"的连续变化过程。由此可见:灌水强度诱导内源激素发生改变,激素特征调控克隆生长格局,克隆生长格局决定种群稳定性。
Studies of the clonal growth regulation of Hippophae rhamnoides ssp. sinensis have been confined to the external mechanisms. This aim of this study was to investigate the response of clonal growth to irrigation intensity and the relationship between the internal mechanism and hormone regulation. With increasing irrigation intensity,the growth capacity,clonal propagation,and clonal expansion of the ramets first increased but then decreased. This trend was also observed for IAA(Indole-3-acetic acid),ZR(Trans-zeatin-riboside),and GA_3(Gibberellic acid),and their ratios to ABA(Abscisic acid),while ABA content showed the opposite trend. The growth capacity,clonal propagation,and clonal expansion were significantly positively correlated with the contents of IAA,ZR,and GA_3,and the ratios of IAA/ABA,ZR/ABA,and GA_3/ABA,while significantly negatively correlated with ABA content. Deficient or excessive irrigation lowered the contentsof IAA,ZR,and GA_3 and the ratios of IAA/ABA,ZR/ABA,and GA_3/ABA and increased ABA content. Clonal growth potential was inhibited and the populations were small ramets,showing a low quantity(sparse distribution) and weak expansion(horizontal root extension and branching). The clone growth was patterned as "guerrilla,"and the population was prone to presenility. Appropriate irrigation intensity increased the contents of IAA,ZR,and GA_3,and their ratios to ABA and decreased the ABA content,indicating the cloning growth potential. The populations were characterized as having large ramets,high quantity(density distribution),and strong expansion ability. The clone growth was patterned as"aggregation,"and populations were stable. With increased irrigation from deficiency to adequacy to excess,the clonal growth was patterned from guerrilla to aggregation to guerrilla in response to the irrigation intensity by regulation of hormone conditions,and the population stability changed from low to high to low. The change in endogenous hormones,which was induced by irrigation intensity,regulated the clonal growth pattern,which determined the population stability.
Studies of the clonal growth regulation of Hippophae rhamnoides ssp. sinensis have been confined to the external mechanisms. This aim of this study was to investigate the response of clonal growth to irrigation intensity and the relationship between the internal mechanism and hormone regulation. With increasing irrigation intensity,the growth capacity,clonal propagation,and clonal expansion of the ramets first increased but then decreased. This trend was also observed for IAA(Indole-3-acetic acid),ZR(Trans-zeatin-riboside),and GA_3(Gibberellic acid),and their ratios to ABA(Abscisic acid),while ABA content showed the opposite trend. The growth capacity,clonal propagation,and clonal expansion were significantly positively correlated with the contents of IAA,ZR,and GA_3,and the ratios of IAA/ABA,ZR/ABA,and GA_3/ABA,while significantly negatively correlated with ABA content. Deficient or excessive irrigation lowered the contentsof IAA,ZR,and GA_3 and the ratios of IAA/ABA,ZR/ABA,and GA_3/ABA and increased ABA content. Clonal growth potential was inhibited and the populations were small ramets,showing a low quantity(sparse distribution) and weak expansion(horizontal root extension and branching). The clone growth was patterned as "guerrilla,"and the population was prone to presenility. Appropriate irrigation intensity increased the contents of IAA,ZR,and GA_3,and their ratios to ABA and decreased the ABA content,indicating the cloning growth potential. The populations were characterized as having large ramets,high quantity(density distribution),and strong expansion ability. The clone growth was patterned as"aggregation,"and populations were stable. With increased irrigation from deficiency to adequacy to excess,the clonal growth was patterned from guerrilla to aggregation to guerrilla in response to the irrigation intensity by regulation of hormone conditions,and the population stability changed from low to high to low. The change in endogenous hormones,which was induced by irrigation intensity,regulated the clonal growth pattern,which determined the population stability.
引文
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[3]董鸣.资源异质性环境中的植物克隆生长:觅食行为.植物学报,1996,38(10):828-835.
[4]Luo D,Qian Y Q,Han L,Liu J X,Sun Z Y.Phenotypic responses of a stoloniferous clonal plant Buchloe dactyloides to scale-dependent nutrient heterogeneity.PLo S One,2013,8(6):e67396.
[5]Yan X,Wang H W,Wang Q F,Rudstam L G.Risk spreading,habitat selection and division of biomass in a submerged clonal plant:responses to heterogeneous copper pollution.Environmental Pollution,2013,174:114-120.
[6]汤俊兵,肖燕,安树青.根茎克隆植物生态学研究进展.生态学报,2010,30(11):3028-3036.
[7]Wang J C,Shi X,Zhang D Y,Yin L K.Phenotypic plasticity in response to soil moisture availability in the clonal plant Eremosparton songoricum(Litv.)Vass.Journal of Arid Land,2011,3(1):34-39.
[8]Wang P,Lei J P,Li M H,Yu F H.Spatial heterogeneity in light supply affects intraspecific competition of a stoloniferous clonal plant.PLo S One,2012,7(6):e39105.
[9]何军,赵聪蛟,清华,甘琳,安树青.土壤水分条件对克隆植物互花米草表型可塑性的影响.生态学报,2009,29(7):3518-3524.
[10]朱万泽,王金锡,罗成荣,段学梅.森林萌生更新研究进展.林业科学,2007,43(9):74-82.
[11]Cline M G.Apical dominance.The Botanical Review,1991,57(4):318-358.
[12]Wan X C,Landhusser S M,Lieffers V J,Zwiazek,J J.Signals controlling root suckering and adventitious shoot formation in aspen(Populus tremuloides).Tree Physiology,2006,26(5):681-687.
[13]Mader J C,Turnbull C G N,Emery R J N.Transport and metabolism of xylem cytokinins during lateral bud release in decapitated chickpea(Cicer arietinum)seedlings.Physiologia Plantarum,2003,117(1):118-129.
[14]许智宏,薛红卫.植物激素作用的分子机理.上海:上海科学技术出版社,2012.
[15]Liu Y Y,Will R E,Tauer C G.Gene level responses of shortleaf pine and loblolly pine to top removal.Tree Genetics&Genomes,2011,7(5):969-986.
[16]高志义,张玉胜.沙棘根系特性的观察与研究.北京林业大学学报,1989,11(4):53-59.
[17]李根前,黄宝龙,唐德瑞,赵一庆,王迪海.毛乌素沙地中国沙棘无性系种群年龄结构动态与遗传后果研究.应用生态学报,2001,12(3):347-350.
[18]李根前,赵粉侠,李秀寨,韦宇.毛乌素沙地中国沙棘种群数量动态研究.林业科学,2004,40(1):180-184.
[19]贺斌,李根前,徐德兵,李甜江,倪金碧.沙棘克隆生长及其生态学意义.西北林学院学报,2006,21(3):54-59.
[20]李根前,黄宝龙,唐德瑞,赵一庆,王迪海.毛乌素沙地中国沙棘无性系生长调节.应用生态学报,2001,12(5):682-686.
[21]郭峰,唐翠平,徐德兵,李根前,贺斌,李甜江.木本克隆植物中国沙棘种群林缘扩散规律研究.云南大学学报:自然科学版,2015,37(2):310-316.
[22]惠兴学,洪新,于欣,张立霞.辽西地区沙棘退化林分的修复与前景.国际沙棘研究与开发,2009,7(4):24-27,31-31.
[23]李秀寨,李根前,韦宇,贺斌.中国沙棘大面积死亡原因的探讨.沙棘,2005,18(1):24-28.
[24]张增悦,姜准,李甜江,肖智勇,李根前.毛乌素沙地中国沙棘人工林早衰原因与特点.西北林学院学报,2016,31(6):1-6.
[25]Cao Z L,Li T J,Li G Q,Liu C H,Gao H Y,Dai G H,Xiao Z Y,Li S L.Modular growth and clonal propagation of Hippophae rhamnoides subsp.sinensis in response to irrigation intensity.Journal of Forestry Research,2016,27(5):1019-1028.
[26]贺斌,赵粉侠,李根前,马先锋,徐德兵,李甜江.毛乌素沙地中国沙棘克隆生长对土壤水分含量的响应.南京林业大学学报:自然科学版,2012,36(4):46-50.
[27]曾诚,陈贝贝,肖智勇,李甜江,李孙玲,代光辉,李根前.毛乌素沙地土壤水分对中国沙棘人工林稳定性及生产力的影响.林业资源管理,2016,(1):99-104.
[28]李甜江,李根前,徐德兵,贺斌,高甲荣.中国沙棘克隆生长对灌水强度的响应.生态学报,2010,30(24):6952-6960.
[29]吴颂如,陈婉芬,周燮.酶联免疫法(ELISA)测定内源植物激素.植物生理学报,1988,24(5):53-57.
[30]潘瑞炽.植物生理学(第四版).北京:高等教育出版社,2001:229-230.
[31]Davies P J.Plant Hormones:Physiology,Biochemistry and Molecular Biology.3rd ed.Dordrecht:Springer,1995.
[32]陈尚,李自珍,王刚.克隆植物生长型的研究进展.生态学杂志,1997,16(4):59-63.
[33]李甜江,李根前,贺斌,徐德兵.克隆植物中国沙棘生长对外源植物激素的响应.生态学报,2010,30(3):659-667.
[34]刘瑞香,杨劼,高丽.中国沙棘和俄罗斯沙棘叶片在不同土壤水分条件下脯氨酸、可溶性糖及内源激素含量的变化.水土保持学报,2005,19(3):148-151,169-169.
[35]李丽霞,梁宗锁,魏宇昆,韩蕊莲.土壤干旱胁迫下沙棘休眠、萌芽期内源激素变化及外源GA3的调节.西北林学院学报,2001,16(2):10-14.
[2]Silvertown J W.Introduction to Plant Population Ecology.2nd ed.Longman,London:John Wiley&Sons,Inc,1987.
[3]董鸣.资源异质性环境中的植物克隆生长:觅食行为.植物学报,1996,38(10):828-835.
[4]Luo D,Qian Y Q,Han L,Liu J X,Sun Z Y.Phenotypic responses of a stoloniferous clonal plant Buchloe dactyloides to scale-dependent nutrient heterogeneity.PLo S One,2013,8(6):e67396.
[5]Yan X,Wang H W,Wang Q F,Rudstam L G.Risk spreading,habitat selection and division of biomass in a submerged clonal plant:responses to heterogeneous copper pollution.Environmental Pollution,2013,174:114-120.
[6]汤俊兵,肖燕,安树青.根茎克隆植物生态学研究进展.生态学报,2010,30(11):3028-3036.
[7]Wang J C,Shi X,Zhang D Y,Yin L K.Phenotypic plasticity in response to soil moisture availability in the clonal plant Eremosparton songoricum(Litv.)Vass.Journal of Arid Land,2011,3(1):34-39.
[8]Wang P,Lei J P,Li M H,Yu F H.Spatial heterogeneity in light supply affects intraspecific competition of a stoloniferous clonal plant.PLo S One,2012,7(6):e39105.
[9]何军,赵聪蛟,清华,甘琳,安树青.土壤水分条件对克隆植物互花米草表型可塑性的影响.生态学报,2009,29(7):3518-3524.
[10]朱万泽,王金锡,罗成荣,段学梅.森林萌生更新研究进展.林业科学,2007,43(9):74-82.
[11]Cline M G.Apical dominance.The Botanical Review,1991,57(4):318-358.
[12]Wan X C,Landhusser S M,Lieffers V J,Zwiazek,J J.Signals controlling root suckering and adventitious shoot formation in aspen(Populus tremuloides).Tree Physiology,2006,26(5):681-687.
[13]Mader J C,Turnbull C G N,Emery R J N.Transport and metabolism of xylem cytokinins during lateral bud release in decapitated chickpea(Cicer arietinum)seedlings.Physiologia Plantarum,2003,117(1):118-129.
[14]许智宏,薛红卫.植物激素作用的分子机理.上海:上海科学技术出版社,2012.
[15]Liu Y Y,Will R E,Tauer C G.Gene level responses of shortleaf pine and loblolly pine to top removal.Tree Genetics&Genomes,2011,7(5):969-986.
[16]高志义,张玉胜.沙棘根系特性的观察与研究.北京林业大学学报,1989,11(4):53-59.
[17]李根前,黄宝龙,唐德瑞,赵一庆,王迪海.毛乌素沙地中国沙棘无性系种群年龄结构动态与遗传后果研究.应用生态学报,2001,12(3):347-350.
[18]李根前,赵粉侠,李秀寨,韦宇.毛乌素沙地中国沙棘种群数量动态研究.林业科学,2004,40(1):180-184.
[19]贺斌,李根前,徐德兵,李甜江,倪金碧.沙棘克隆生长及其生态学意义.西北林学院学报,2006,21(3):54-59.
[20]李根前,黄宝龙,唐德瑞,赵一庆,王迪海.毛乌素沙地中国沙棘无性系生长调节.应用生态学报,2001,12(5):682-686.
[21]郭峰,唐翠平,徐德兵,李根前,贺斌,李甜江.木本克隆植物中国沙棘种群林缘扩散规律研究.云南大学学报:自然科学版,2015,37(2):310-316.
[22]惠兴学,洪新,于欣,张立霞.辽西地区沙棘退化林分的修复与前景.国际沙棘研究与开发,2009,7(4):24-27,31-31.
[23]李秀寨,李根前,韦宇,贺斌.中国沙棘大面积死亡原因的探讨.沙棘,2005,18(1):24-28.
[24]张增悦,姜准,李甜江,肖智勇,李根前.毛乌素沙地中国沙棘人工林早衰原因与特点.西北林学院学报,2016,31(6):1-6.
[25]Cao Z L,Li T J,Li G Q,Liu C H,Gao H Y,Dai G H,Xiao Z Y,Li S L.Modular growth and clonal propagation of Hippophae rhamnoides subsp.sinensis in response to irrigation intensity.Journal of Forestry Research,2016,27(5):1019-1028.
[26]贺斌,赵粉侠,李根前,马先锋,徐德兵,李甜江.毛乌素沙地中国沙棘克隆生长对土壤水分含量的响应.南京林业大学学报:自然科学版,2012,36(4):46-50.
[27]曾诚,陈贝贝,肖智勇,李甜江,李孙玲,代光辉,李根前.毛乌素沙地土壤水分对中国沙棘人工林稳定性及生产力的影响.林业资源管理,2016,(1):99-104.
[28]李甜江,李根前,徐德兵,贺斌,高甲荣.中国沙棘克隆生长对灌水强度的响应.生态学报,2010,30(24):6952-6960.
[29]吴颂如,陈婉芬,周燮.酶联免疫法(ELISA)测定内源植物激素.植物生理学报,1988,24(5):53-57.
[30]潘瑞炽.植物生理学(第四版).北京:高等教育出版社,2001:229-230.
[31]Davies P J.Plant Hormones:Physiology,Biochemistry and Molecular Biology.3rd ed.Dordrecht:Springer,1995.
[32]陈尚,李自珍,王刚.克隆植物生长型的研究进展.生态学杂志,1997,16(4):59-63.
[33]李甜江,李根前,贺斌,徐德兵.克隆植物中国沙棘生长对外源植物激素的响应.生态学报,2010,30(3):659-667.
[34]刘瑞香,杨劼,高丽.中国沙棘和俄罗斯沙棘叶片在不同土壤水分条件下脯氨酸、可溶性糖及内源激素含量的变化.水土保持学报,2005,19(3):148-151,169-169.
[35]李丽霞,梁宗锁,魏宇昆,韩蕊莲.土壤干旱胁迫下沙棘休眠、萌芽期内源激素变化及外源GA3的调节.西北林学院学报,2001,16(2):10-14.