红松种子园种群表型多样性研究
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摘要
为揭示红松(Pinus koraiensis Sieb. et Zucc.)不同种群的表型分化程度和变异规律,以吉林省露水河红松种子园6个种群红松为研究对象,采用巢式方差分析、多重比较、变异系数和聚类分析等方法对红松的叶片、球果和种子共15个表型性状进行了系统分析。结果表明:(1)红松15个表型性状在种群间和种群内均存在着极显著的差异,红松种群遗变异比较丰富,在松属植物中属于中等水平,其中纬度最低的露水河种群在其中10个表型性状均值上表现出最高值;(2)红松种群间的表型分化系数(Vst)均值为12.39%,种群内的变异(87.61%)大于种群间的变异(12.39%),种群内的变异是主要变异来源;(3)各表型性状平均变异系数为13.28%,变幅为6.16%—31.48%,叶片、球果、种子的平均变异系数依次为:球果17.86%>针叶11.19%>种子9.87%,种子性状最小,成为最稳定的表型性状,带岭和丰林种群表型性状遗传多样性要高于其他种群;(4)利用欧氏平均距离对红松种群进行UPGMA聚类分析,红松种群的表型性状按照地理距离而聚类,可将红松种子园6个种群分为4类,其表型性状跟地形(东北山脉)有一定的契合。红松种群具有中等水平的表型遗传多样性,种群间和种群内均存在丰富的表型变异,研究结果为顺利开展红松种质资源收集、保存,以及良种选育等工作提供依据。
Pinus koraiensis is a rare and protected species in China. In this study, our objectives were to determine the phenotypic variation, distribution mechanisms and patterns of different P. koraiensis populations in a seed orchard. We studied morphological variation among and within six populations of P. koraiensis based on analysis of 15 phenotypic traits(including needle, cone, and seed traits). Nested analysis of variance, coefficient of variation, multi-comparison, and the unweighted pair-group method with arithmetic means(UPGMA) cluster analysis were used to analyse the data. The results showed that:(1) there were significant differences in all 15 traits among and within populations of P. koraiensis, revealing a large variation. Within the genus Pinus, P. koraiensis shows intermediate levels of variation. The most southern population showed the highest mean value in 10 of the 15 traits;(2) mean phenotypic differentiation coefficient(Vst) within populations was 87.61%, higher than that among population(12.39%), indicating that the largest part of phenotypic variation is retained within populations in P. koraiensis;(3) the average value of coefficient of variation(CV) was 13.28%, and ranged from 6.16% to 31.48% among traits. Among traits, the CV decreased from cone(17.86%) to needle(11.19%) to seed traits(9.87%), which were the most stable traits. The phenotypic diversity of DL and FL populations was higher than that of other populations;(4) according to UPGMA cluster analysis, the phenotypic traits of P. koraiensis populations clustered by geographical distance. The six populations P. koraiensis used in this study can be divided into four groups, which are significantly related in topography(mountains in northeast of China). Overall, we observed abundant phenotypic variation among and within populations, which provides critical information for resource collection, conservation, and breeding of this ecologically important species.
Pinus koraiensis is a rare and protected species in China. In this study, our objectives were to determine the phenotypic variation, distribution mechanisms and patterns of different P. koraiensis populations in a seed orchard. We studied morphological variation among and within six populations of P. koraiensis based on analysis of 15 phenotypic traits(including needle, cone, and seed traits). Nested analysis of variance, coefficient of variation, multi-comparison, and the unweighted pair-group method with arithmetic means(UPGMA) cluster analysis were used to analyse the data. The results showed that:(1) there were significant differences in all 15 traits among and within populations of P. koraiensis, revealing a large variation. Within the genus Pinus, P. koraiensis shows intermediate levels of variation. The most southern population showed the highest mean value in 10 of the 15 traits;(2) mean phenotypic differentiation coefficient(Vst) within populations was 87.61%, higher than that among population(12.39%), indicating that the largest part of phenotypic variation is retained within populations in P. koraiensis;(3) the average value of coefficient of variation(CV) was 13.28%, and ranged from 6.16% to 31.48% among traits. Among traits, the CV decreased from cone(17.86%) to needle(11.19%) to seed traits(9.87%), which were the most stable traits. The phenotypic diversity of DL and FL populations was higher than that of other populations;(4) according to UPGMA cluster analysis, the phenotypic traits of P. koraiensis populations clustered by geographical distance. The six populations P. koraiensis used in this study can be divided into four groups, which are significantly related in topography(mountains in northeast of China). Overall, we observed abundant phenotypic variation among and within populations, which provides critical information for resource collection, conservation, and breeding of this ecologically important species.
引文
[1] 顾万春.统计遗传学.北京:科学出版社,2004.
[2] Pigliucci M,Murren C J,Schlichting C D.Phenotypic plasticity and evolution by genetic assimilation.Journal of Experimental Biology,2006,209(12):2362- 2367.
[3] 李斌,顾万春,卢宝明.白皮松天然群体种实性状表型多样性研究.生物多样性,2002,10(2):181- 188.
[4] 李斌,顾万春.松属植物遗传多样性研究进展.遗传,2003,25(6):740- 748.
[5] 李因刚,柳新红,马俊伟,石从广,朱光权.浙江楠种群表型变异.植物生态学报,2014,38(12):1315- 1324.
[6] 李长喜.林木天然群体表型变异研究概述.林业科学研究,1988,1(6):657- 664.
[7] 李帅锋,苏建荣,刘万德,郎学东,张志钧,苏磊,贾呈鑫卓,杨华景.思茅松天然群体种实表型变异.植物生态学报,2013,37(11):998- 1009.
[8] 张翠琴,姬志峰,林丽丽,赵瑞华,王祎玲.五角枫种群表型多样性.生态学报,2015,35(16):5343- 5352.
[9] Xu Y L,Woeste K,Cai N H,Kang X Y,Li G Q,Chen S,Duan A A.Variation in needle and cone traits in natural populations of Pinus yunnanensis.Journal of Forestry Research,2016,27(1):41- 49.
[10] Li Y G,Liu X H,Ma J W,Zhang X M,Xu L A.Phenotypic variation in Phoebe bournei populations preserved in the primary distribution area.Journal of Forestry Research,2018,29(1):35- 44.
[11] Kooke R,Johannes F,Wardenaar R,Becker F,Etcheverry M,Colot V,Vreugdenhil D,Keurentjes J J B.Epigenetic basis of morphological variation and phenotypic plasticity in Arabidopsis thaliana.The Plant Cell,2015,27(2):337- 348.
[12] 傅俊卿.东北天然红松林资源现状与保护经营对策.东北林业大学学报,2009,37(2):75- 78.
[13] 于大炮,周旺明,包也,齐麟,周莉,代力民.天保工程实施以来东北阔叶红松林的可持续经营.生态学报,2015,35(1):10- 17.
[14] 傅立国.中国植物红皮书.北京:科学出版社,1991.
[15] 杨建国,王元兴,王永凡,周春艳,王焕章.红松优良种源的选择.吉林林业科技,1998,5(5):12- 14.
[16] Sui X,Feng F J,Zhao D,Xing M,Sun X Y,Han S J,Li M H.Mating system patterns of natural populations of Pinus koraiensis along its post-glacial colonization route in northeastern China.Genetics and Molecular Research,2015,14(2):4113- 4124.
[17] Feng F J,Han S J,Wang H M.Genetic diversity and genetic differentiation of natural Pinus koraiensis population.Journal of Forestry Research,2006,17(1):21- 24.
[18] Potenko V V,Velikov A V.Genetic diversity and differentiation of natural populations of Pinus koraiensis (SIEB.et Zucc.) in Russia.Silvae Genetica,1998,47(4):202- 208.
[19] 葛颂,王明庥,陈岳武.用同工酶研究马尾松群体的遗传结构.林业科学,1988,24(4):399- 409.
[20] 罗建勋,顾万春.云杉天然群体表型多样性研究.林业科学,2005,41(2):66- 73.
[21] Zas R,Sampedro L.Heritability of seed weight in Maritime pine,a relevant trait in the transmission of environmental maternal effects.Heredity,2015,114(1):116- 124.
[22] Jasińska A K,Boratyńska K,Dering M,Sobierajska K I,Ok T,Romo A,Boratyński A.Distance between south-European and south-west Asiatic refugial areas involved morphological differentiation:Pinus sylvestris case study.Plant Systematics and Evolution,2014,300(6):1487- 1502.
[23] 张振.红松无性系种子形态及营养成分变异研究[D].哈尔滨:东北林业大学,2015.
[24] 陈常美.东北林区主要珍贵用材树种种质遗传多样性评价与保护策略[D].哈尔滨:东北林业大学,2013.
[25] 冯富娟,隋心,张冬东.不同种源红松遗传多样性的研究.林业科技,2008,33(1):1- 4.
[26] Yeh F C,El-Kassaby Y A.Enzyme variation in natural populations of Sitka spruce (Picea sitchensis).1.Genetic variation patterns among trees from 10 IUFRO provenances.Canadian Journal of Forest Research,1980,10(3):415- 422.
[27] El-Kassaby Y A,Sziklai O.Genetic variation of allozyme and quantitative traits in a selected douglas-fir [Pseudotsuga menziesii var.menziesii (Mirb.) Franco] population.Forest Ecology and Management,1982,4(2):115- 126.
[28] 姬明飞,张晓玮,韩瑾,丁东粮.油松天然群体的种实性状表型多样性研究.西北植物学报,2013,33(9):1898- 1905.
[29] Hamrick J L,Godt M J W,Sherman-Broyles S L.Factors influencing levels of genetic diversity in woody plant species.New Forests,1992,6(1/4):95- 124.
[30] 葛颂.同工酶与林木群体遗传变异研究.南京林业大学学报,1988,1(1):68- 77.
[31] 邢猛.天然红松种群交配系统变化规律的研究[D].哈尔滨:东北林业大学,2013.
[32] Hartl D L,Clark A G.Principles of Population Genetics.Sunderland,USA:Sinauer Associates,Inc,1997.
[33] 杜荣骞.生物统计学(第三版).北京:高等教育出版社,2009.
[34] Sorensen F C,Miles R S.Cone and seed weight relationships in douglas-fir from western and central oregon.Ecology,1978,59(4):641- 644.
[35] 童冉,吴小龙,姜丽娜,司倩倩,臧德奎.野生玫瑰种群表型变异.生态学报,2017,37(11):3706- 3715.
[36] 辜云杰,罗建勋,吴远伟,曹小军.川西云杉天然种群表型多样性.植物生态学报,2009,33(2):291- 301.
[37] 张巍,王清君,郭兴.红松不同种源的遗传多样性分析.森林工程,2017,33(2):17- 21.
[2] Pigliucci M,Murren C J,Schlichting C D.Phenotypic plasticity and evolution by genetic assimilation.Journal of Experimental Biology,2006,209(12):2362- 2367.
[3] 李斌,顾万春,卢宝明.白皮松天然群体种实性状表型多样性研究.生物多样性,2002,10(2):181- 188.
[4] 李斌,顾万春.松属植物遗传多样性研究进展.遗传,2003,25(6):740- 748.
[5] 李因刚,柳新红,马俊伟,石从广,朱光权.浙江楠种群表型变异.植物生态学报,2014,38(12):1315- 1324.
[6] 李长喜.林木天然群体表型变异研究概述.林业科学研究,1988,1(6):657- 664.
[7] 李帅锋,苏建荣,刘万德,郎学东,张志钧,苏磊,贾呈鑫卓,杨华景.思茅松天然群体种实表型变异.植物生态学报,2013,37(11):998- 1009.
[8] 张翠琴,姬志峰,林丽丽,赵瑞华,王祎玲.五角枫种群表型多样性.生态学报,2015,35(16):5343- 5352.
[9] Xu Y L,Woeste K,Cai N H,Kang X Y,Li G Q,Chen S,Duan A A.Variation in needle and cone traits in natural populations of Pinus yunnanensis.Journal of Forestry Research,2016,27(1):41- 49.
[10] Li Y G,Liu X H,Ma J W,Zhang X M,Xu L A.Phenotypic variation in Phoebe bournei populations preserved in the primary distribution area.Journal of Forestry Research,2018,29(1):35- 44.
[11] Kooke R,Johannes F,Wardenaar R,Becker F,Etcheverry M,Colot V,Vreugdenhil D,Keurentjes J J B.Epigenetic basis of morphological variation and phenotypic plasticity in Arabidopsis thaliana.The Plant Cell,2015,27(2):337- 348.
[12] 傅俊卿.东北天然红松林资源现状与保护经营对策.东北林业大学学报,2009,37(2):75- 78.
[13] 于大炮,周旺明,包也,齐麟,周莉,代力民.天保工程实施以来东北阔叶红松林的可持续经营.生态学报,2015,35(1):10- 17.
[14] 傅立国.中国植物红皮书.北京:科学出版社,1991.
[15] 杨建国,王元兴,王永凡,周春艳,王焕章.红松优良种源的选择.吉林林业科技,1998,5(5):12- 14.
[16] Sui X,Feng F J,Zhao D,Xing M,Sun X Y,Han S J,Li M H.Mating system patterns of natural populations of Pinus koraiensis along its post-glacial colonization route in northeastern China.Genetics and Molecular Research,2015,14(2):4113- 4124.
[17] Feng F J,Han S J,Wang H M.Genetic diversity and genetic differentiation of natural Pinus koraiensis population.Journal of Forestry Research,2006,17(1):21- 24.
[18] Potenko V V,Velikov A V.Genetic diversity and differentiation of natural populations of Pinus koraiensis (SIEB.et Zucc.) in Russia.Silvae Genetica,1998,47(4):202- 208.
[19] 葛颂,王明庥,陈岳武.用同工酶研究马尾松群体的遗传结构.林业科学,1988,24(4):399- 409.
[20] 罗建勋,顾万春.云杉天然群体表型多样性研究.林业科学,2005,41(2):66- 73.
[21] Zas R,Sampedro L.Heritability of seed weight in Maritime pine,a relevant trait in the transmission of environmental maternal effects.Heredity,2015,114(1):116- 124.
[22] Jasińska A K,Boratyńska K,Dering M,Sobierajska K I,Ok T,Romo A,Boratyński A.Distance between south-European and south-west Asiatic refugial areas involved morphological differentiation:Pinus sylvestris case study.Plant Systematics and Evolution,2014,300(6):1487- 1502.
[23] 张振.红松无性系种子形态及营养成分变异研究[D].哈尔滨:东北林业大学,2015.
[24] 陈常美.东北林区主要珍贵用材树种种质遗传多样性评价与保护策略[D].哈尔滨:东北林业大学,2013.
[25] 冯富娟,隋心,张冬东.不同种源红松遗传多样性的研究.林业科技,2008,33(1):1- 4.
[26] Yeh F C,El-Kassaby Y A.Enzyme variation in natural populations of Sitka spruce (Picea sitchensis).1.Genetic variation patterns among trees from 10 IUFRO provenances.Canadian Journal of Forest Research,1980,10(3):415- 422.
[27] El-Kassaby Y A,Sziklai O.Genetic variation of allozyme and quantitative traits in a selected douglas-fir [Pseudotsuga menziesii var.menziesii (Mirb.) Franco] population.Forest Ecology and Management,1982,4(2):115- 126.
[28] 姬明飞,张晓玮,韩瑾,丁东粮.油松天然群体的种实性状表型多样性研究.西北植物学报,2013,33(9):1898- 1905.
[29] Hamrick J L,Godt M J W,Sherman-Broyles S L.Factors influencing levels of genetic diversity in woody plant species.New Forests,1992,6(1/4):95- 124.
[30] 葛颂.同工酶与林木群体遗传变异研究.南京林业大学学报,1988,1(1):68- 77.
[31] 邢猛.天然红松种群交配系统变化规律的研究[D].哈尔滨:东北林业大学,2013.
[32] Hartl D L,Clark A G.Principles of Population Genetics.Sunderland,USA:Sinauer Associates,Inc,1997.
[33] 杜荣骞.生物统计学(第三版).北京:高等教育出版社,2009.
[34] Sorensen F C,Miles R S.Cone and seed weight relationships in douglas-fir from western and central oregon.Ecology,1978,59(4):641- 644.
[35] 童冉,吴小龙,姜丽娜,司倩倩,臧德奎.野生玫瑰种群表型变异.生态学报,2017,37(11):3706- 3715.
[36] 辜云杰,罗建勋,吴远伟,曹小军.川西云杉天然种群表型多样性.植物生态学报,2009,33(2):291- 301.
[37] 张巍,王清君,郭兴.红松不同种源的遗传多样性分析.森林工程,2017,33(2):17- 21.
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