尾叶桉与巨桉杂种F_1代生长性状遗传分析
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摘要
【目的】测定尾叶桉与巨桉杂种F_1多年度的生长性状,估算其遗传参数,开展优良杂交组合及杂种优良单株选择,为桉树亲本选配和杂种应用提供科学依据。【方法】以尾叶桉与巨桉6×6析因交配F_1代杂种为材料,应用ASReml-R软件估算1、2、4.5、8.3和10.3年生时生长性状的遗传力、显性效应、加性效应、亲本效应和配合力等遗传参数;计算每个杂交组合单株材积的平均值,以所有组合的平均值+标准差为标准进行优良杂交组合选择;基于所有单株的材积,采用平均值+2倍标准差(标准Ⅰ)和平均值+3倍标准差(标准Ⅱ)2个标准进行杂种优良单株选择。【结果】尾巨桉F_1代杂种在各年龄时的生长表现均优于对照尾叶桉亲本自由授粉子代。每个年份树高、胸径和单株材积在组合间均存在极显著(P<0.01)或显著(P<0.05)差异,在区组间差异不显著。1年生和2年生时杂种各生长性状的母本加性方差分量低于父本加性方差分量, 4.5年生时父、母本加性方差分量较为接近,8.3和10.3年生时母本加性方差分量则高于父本加性方差分量。随年龄的增长,杂种的树高和胸径单株遗传力呈现由中度至低度遗传控制的变化趋势,杂种家系遗传力呈现高—中—低度遗传控制的趋势,且杂种家系遗传力大于单株遗传力。显性方差与加性方差的比值则随年龄的增长呈递增趋势。年-年和性状-性状双变量遗传、表型呈显著或极显著正相关。4.5年生时,母本U21及父本G8的一般杂交力(GHA)最高,U21×G5组合的特殊杂交力(SHA)最高; 8.3年生时,母本U21及父本G19的GHA最高,U2×G19的SHA最高。【结论】尾巨桉F_1杂种1～2年生时父本效应大于母本效应,且加性效应大于显性效应,4.5年生时父、母本效应对杂种生长的贡献率相当,8.3～10.3年生时杂种的母本效应大于父本效应,且显性效应大于加性效应。同时评选出7个优良杂种,筛选出1个优良母本和2个优良父本,并依据标准Ⅰ、Ⅱ分别选出40株和16株优树,为今后大规模人工制种及推广应用提供可靠的亲本材料和优良无性系。
【Objective】 Multiple-year growth traits of Eucalyptus urophylla×E. grandis F_1 hybrids were investigated, their genetic parameters were estimated, and superior hybrids and plus trees were selected aiming to provide a scientific basis for selection of mating parents and their hybrids.【Method】 The F_1 hybrids of E. urophylla×E. grandis from factorial mating system were studied using the open-pollinated progenies of E. urophylla as controls. ASReml-R software was applied to assess heritability, dominance effect,additive effect, parental effect and combining abilities of growth traits when 1, 2, 4.5, 8.3 and 10.3 years old. Mean single-tree volume was calculated for each hybrid combination, and superior combinations were determined as those with volume above mean+standard deviation of all combinations. Selection of plus trees was conducted with two standards, I: mean + two times standard deviation, and II: mean + three times standard deviation.【Result】 Growth performance of hybrids was obviously better than that of open-pollinated maternal off-springs at each age. Tree height, diameter at breast height(DBH) and individual volume at each age were significantly(P<0.05 or P<0.01) different among hybrid combinations, while no significant difference was observed among blocks. Maternal variance component of each growth trait was lower than paternal variance component when one or two years old, they were equivalent to each other when 4.5 years old, while maternal variance component was higher than paternal variance component when 8.3 or 10.3 years old. With age increase, single-tree heritability(h~2) generally tended to vary from middle to low, family heritability(h■) from high to middle to low, while additive-to-dominance variance ratio(σ~2_D/σ~2_A) increased for tree height and DBH. Year-year and trait-trait genetic and phenotypic correlations were significant among hybrids. The general hybridizing abilities(GHA) of U21 and G8 were the highest among maternal and paternal parents, respectively, and the specific hybridizing ability(SHA) of U21×G5 was the highest among all cross combinations when 4.5 years old. When 8.3 years old, GHA of U21 and G19 were the highest, and SHA of U2×G19 was the highest.【Conclusion】 Maternal and dominance effect of each growth trait was lower than paternal and additive effect when one or two years old, respectively,maternal and paternal effect were equivalent to each other when 4.5 years old, while maternal and dominance effect was higher than paternal and additive effect when 8.3 or 10.3 years old. A total of 7 superior cross combinations, one superior maternal parent and two paternal parents were selected, and 40 and 16 elite individuals were determined according to Standards Ⅰ and Ⅱ, respectively. The findings can provide reliable parent materials for a large number of artificial seed production and clones for eucalypt extension in the future.
【Objective】 Multiple-year growth traits of Eucalyptus urophylla×E. grandis F_1 hybrids were investigated, their genetic parameters were estimated, and superior hybrids and plus trees were selected aiming to provide a scientific basis for selection of mating parents and their hybrids.【Method】 The F_1 hybrids of E. urophylla×E. grandis from factorial mating system were studied using the open-pollinated progenies of E. urophylla as controls. ASReml-R software was applied to assess heritability, dominance effect,additive effect, parental effect and combining abilities of growth traits when 1, 2, 4.5, 8.3 and 10.3 years old. Mean single-tree volume was calculated for each hybrid combination, and superior combinations were determined as those with volume above mean+standard deviation of all combinations. Selection of plus trees was conducted with two standards, I: mean + two times standard deviation, and II: mean + three times standard deviation.【Result】 Growth performance of hybrids was obviously better than that of open-pollinated maternal off-springs at each age. Tree height, diameter at breast height(DBH) and individual volume at each age were significantly(P<0.05 or P<0.01) different among hybrid combinations, while no significant difference was observed among blocks. Maternal variance component of each growth trait was lower than paternal variance component when one or two years old, they were equivalent to each other when 4.5 years old, while maternal variance component was higher than paternal variance component when 8.3 or 10.3 years old. With age increase, single-tree heritability(h~2) generally tended to vary from middle to low, family heritability(h■) from high to middle to low, while additive-to-dominance variance ratio(σ~2_D/σ~2_A) increased for tree height and DBH. Year-year and trait-trait genetic and phenotypic correlations were significant among hybrids. The general hybridizing abilities(GHA) of U21 and G8 were the highest among maternal and paternal parents, respectively, and the specific hybridizing ability(SHA) of U21×G5 was the highest among all cross combinations when 4.5 years old. When 8.3 years old, GHA of U21 and G19 were the highest, and SHA of U2×G19 was the highest.【Conclusion】 Maternal and dominance effect of each growth trait was lower than paternal and additive effect when one or two years old, respectively,maternal and paternal effect were equivalent to each other when 4.5 years old, while maternal and dominance effect was higher than paternal and additive effect when 8.3 or 10.3 years old. A total of 7 superior cross combinations, one superior maternal parent and two paternal parents were selected, and 40 and 16 elite individuals were determined according to Standards Ⅰ and Ⅱ, respectively. The findings can provide reliable parent materials for a large number of artificial seed production and clones for eucalypt extension in the future.
引文
鲍甫成,罗建举.2002.尾叶桉×巨桉F1子代生长和材质性状遗传变异性研究.林业科学,38(3):124-130.(Bao F C,Luo J J.2002.Genetic variation in growth and wood properties of F1 progenies of Eucalyptus urophylla×E.grandis.Scientia Silvae Sinicae,38(3):124-130.[in Chinese])
陈永忠,王德斌,彭邵锋,等.2008.油茶优良杂交组合选育研究.中南林业科技大学学报,28(4):27-35.(Chen Y Z,Wang D B,Peng S F,et al.2008.Selective breeding of elite crossbred siblings of Camellia oleifera.Journal of Central South University of Forestry & Technology,28(4):27-35.[in Chinese])
何旭东,李发根,翁启杰,等.2010.尾叶桉×细叶桉杂种生长与耐寒性的联合选择.中南林业科技大学学报,30(8):68-70.(He X D,Li F G,Weng Q J,et al.2010.Selection of Eucalptus urophylla ×E.tereticornis hybrids in growth and cold hardiness.Journal of Central South University of Forestry & Technology,30(8):68-70.[in Chinese])
Jacobs M R.1979.桉树栽培.联合国粮农组织,624-625.(Jacobs M R.1979.Eucalyptus for Planting.Food and Agriculture Organization of the United Nations,624-625.[in Chinese])
Pegg Richard,李海佳,李宏伟,等.2006.广西桉树人工杂交育种研究.广西林业科学,35(4):238-242.(Pegg Richard,Li H J,Li H W,et al.2006.Cross breeding of Eucalyptus in Guangxi.Guangxi Forestry Science,35(4):238-242.[in Chinese])
谭小梅,金国庆,邵纪清,等.2011.马尾松巢式交配子代生长和木材基本密度遗传分析.林业科学,47(6):30-35.(Tang X M,Jin G Q,Shao J Q,et al.2011.Genetic analysis of growth and wood density of progeny of nested mating Pinus massoniana.Scientia Silvae Sinicae,47(6):30-35.[in Chinese])
王豁然.2010.桉树生物学概论.北京:科学出版社,65-68.(Wang H R.2010.A Chinese appreciation of Eucalypts.Beijing:Science Press,65-68.[in Chinese])
翁启杰,赖秋香,李发根,等.2015.尾叶桉×邓恩桉早期生长和耐寒性的遗传分析.南京林业大学学报,39(5):33-38.(Weng Q J,Lai Q X,Li F G,et al.2015.Genetic analysis on early growth and cold tolerance of Eucalyptus urophylla × E.dunnii hybrids.Journal of Nanjing Forestry University,39(5):33-38.[in Chinese])
赵奋成,李宪政,张应中,等.2006.湿地松与洪都拉斯加勒比松的杂交效果分析.林业科学研究,19(4):409-415.(Zhao F C,Li X Z,Zhang Y Z,et al.2006.Hybridizing effect analysis between Pinus elliottii var.elliottii and Pinus caribaea var.hondurensis.Forest Research,19(4):409-415.[in Chinese])
Bison O,Ramalho M A P,Resende G D S P,et al.2006.Comparison between open pollinated progenies and hybrids performance in Eucalyptus grandis and Eucalyptus urophylla.Silvae Genetica,55(4/5):192-196.
Bison O,Ramalho M A P,Resende G D S P.2007.Combining ability of elite clones of Eucalyptus grandis and Eucalyptus urophylla with Eucalyptus globulus.Genetic Molecular Biology,30(2):417-422.
Bouvet J M,Vigneron P.1996.Genetic structure of Eucalyptus urophylla× E.grandis population in the reciprocal recurrent selection scheme in the Congo// Dieters M J,Matheson A C,Nikles D G,et al.Tree improvement for sustainable tropical forestry.QFRI,Brisbane,Caloundra,Queensland,127-132.
Bouvet J M,Saya A,Vigneron P.2009.Trends in additive,dominance and environmental effects with age for growth traits in Eucalyptus hybrid populations.Euphytica,165(1):35-54.
Campinhos E Jr,Ikemori Y K.1989.Selection and management of the basic population Eucalyptus grandis and E.urophylla established at Aracruz for the long term breeding programme//Gibson G I,Griffin A R,Matheson A C.Breeding tropical trees:population structure and genetic improvement strategies in clonal and seedling forestry.Proceedings of IUFRO conference,Pattaya,Thailand.Oxford Forestry Institute,Oxford,169–175.
Gilmour A R,Gogel B J,Cullis B R,et al.2009.ASReml user guide release 3.0.Hemel Hempstead:VSN International Ltd.
Gwaze D P,Bridgwater F E,Lowe W J.2000.Performance of interspecific F1 eucalypt hybrids in Zimbabwe.Forest Genetics,7(4):295-303.
Madhibha T,Murepa R,Musokonyi C,et al.2013.Genetic parameter estimates for interspecific Eucalyptus hybrids and implications for hybrid breeding strategy.New Forests,4(1):63-84.
Paul A D,Foster G S,Caldwell T,et al.1997.Trends in genetic and environmental parameters for height,diameter,and volume in a multilocation clonal study with Loblolly pine.Forest Science,43(1):87-98.
Retief E C L,Stanger T K.2009.Genetic parameters of pure and hybrid populations of Eucalyptus grandis and E.urophylla and implications for hybrid breeding strategy.Journal of the South African Forestry Association,71(2):133-140.
Shi T,Arnold R J,Kang W,et al.2016.Genetic variation and gains for two generations of Eucalyptus dunnii in China.Australian Forestry,79(1):15-24.
Volker P W.2008.Genetic parameters of intra- and inter-specific hybrids of Eucalyptus globulus and E.nitens.Tree Genetics & Genomes,4(3):445-460.
Zhu Y,Wu S,Xu J,et al.2017.Genetic parameters for growth traits and stem-straightness in Eucalyptus urophylla × E.camaldulensis hybrids from a reciprocal mating design.Euphytica,213(7):142-158.
陈永忠,王德斌,彭邵锋,等.2008.油茶优良杂交组合选育研究.中南林业科技大学学报,28(4):27-35.(Chen Y Z,Wang D B,Peng S F,et al.2008.Selective breeding of elite crossbred siblings of Camellia oleifera.Journal of Central South University of Forestry & Technology,28(4):27-35.[in Chinese])
何旭东,李发根,翁启杰,等.2010.尾叶桉×细叶桉杂种生长与耐寒性的联合选择.中南林业科技大学学报,30(8):68-70.(He X D,Li F G,Weng Q J,et al.2010.Selection of Eucalptus urophylla ×E.tereticornis hybrids in growth and cold hardiness.Journal of Central South University of Forestry & Technology,30(8):68-70.[in Chinese])
Jacobs M R.1979.桉树栽培.联合国粮农组织,624-625.(Jacobs M R.1979.Eucalyptus for Planting.Food and Agriculture Organization of the United Nations,624-625.[in Chinese])
Pegg Richard,李海佳,李宏伟,等.2006.广西桉树人工杂交育种研究.广西林业科学,35(4):238-242.(Pegg Richard,Li H J,Li H W,et al.2006.Cross breeding of Eucalyptus in Guangxi.Guangxi Forestry Science,35(4):238-242.[in Chinese])
谭小梅,金国庆,邵纪清,等.2011.马尾松巢式交配子代生长和木材基本密度遗传分析.林业科学,47(6):30-35.(Tang X M,Jin G Q,Shao J Q,et al.2011.Genetic analysis of growth and wood density of progeny of nested mating Pinus massoniana.Scientia Silvae Sinicae,47(6):30-35.[in Chinese])
王豁然.2010.桉树生物学概论.北京:科学出版社,65-68.(Wang H R.2010.A Chinese appreciation of Eucalypts.Beijing:Science Press,65-68.[in Chinese])
翁启杰,赖秋香,李发根,等.2015.尾叶桉×邓恩桉早期生长和耐寒性的遗传分析.南京林业大学学报,39(5):33-38.(Weng Q J,Lai Q X,Li F G,et al.2015.Genetic analysis on early growth and cold tolerance of Eucalyptus urophylla × E.dunnii hybrids.Journal of Nanjing Forestry University,39(5):33-38.[in Chinese])
赵奋成,李宪政,张应中,等.2006.湿地松与洪都拉斯加勒比松的杂交效果分析.林业科学研究,19(4):409-415.(Zhao F C,Li X Z,Zhang Y Z,et al.2006.Hybridizing effect analysis between Pinus elliottii var.elliottii and Pinus caribaea var.hondurensis.Forest Research,19(4):409-415.[in Chinese])
Bison O,Ramalho M A P,Resende G D S P,et al.2006.Comparison between open pollinated progenies and hybrids performance in Eucalyptus grandis and Eucalyptus urophylla.Silvae Genetica,55(4/5):192-196.
Bison O,Ramalho M A P,Resende G D S P.2007.Combining ability of elite clones of Eucalyptus grandis and Eucalyptus urophylla with Eucalyptus globulus.Genetic Molecular Biology,30(2):417-422.
Bouvet J M,Vigneron P.1996.Genetic structure of Eucalyptus urophylla× E.grandis population in the reciprocal recurrent selection scheme in the Congo// Dieters M J,Matheson A C,Nikles D G,et al.Tree improvement for sustainable tropical forestry.QFRI,Brisbane,Caloundra,Queensland,127-132.
Bouvet J M,Saya A,Vigneron P.2009.Trends in additive,dominance and environmental effects with age for growth traits in Eucalyptus hybrid populations.Euphytica,165(1):35-54.
Campinhos E Jr,Ikemori Y K.1989.Selection and management of the basic population Eucalyptus grandis and E.urophylla established at Aracruz for the long term breeding programme//Gibson G I,Griffin A R,Matheson A C.Breeding tropical trees:population structure and genetic improvement strategies in clonal and seedling forestry.Proceedings of IUFRO conference,Pattaya,Thailand.Oxford Forestry Institute,Oxford,169–175.
Gilmour A R,Gogel B J,Cullis B R,et al.2009.ASReml user guide release 3.0.Hemel Hempstead:VSN International Ltd.
Gwaze D P,Bridgwater F E,Lowe W J.2000.Performance of interspecific F1 eucalypt hybrids in Zimbabwe.Forest Genetics,7(4):295-303.
Madhibha T,Murepa R,Musokonyi C,et al.2013.Genetic parameter estimates for interspecific Eucalyptus hybrids and implications for hybrid breeding strategy.New Forests,4(1):63-84.
Paul A D,Foster G S,Caldwell T,et al.1997.Trends in genetic and environmental parameters for height,diameter,and volume in a multilocation clonal study with Loblolly pine.Forest Science,43(1):87-98.
Retief E C L,Stanger T K.2009.Genetic parameters of pure and hybrid populations of Eucalyptus grandis and E.urophylla and implications for hybrid breeding strategy.Journal of the South African Forestry Association,71(2):133-140.
Shi T,Arnold R J,Kang W,et al.2016.Genetic variation and gains for two generations of Eucalyptus dunnii in China.Australian Forestry,79(1):15-24.
Volker P W.2008.Genetic parameters of intra- and inter-specific hybrids of Eucalyptus globulus and E.nitens.Tree Genetics & Genomes,4(3):445-460.
Zhu Y,Wu S,Xu J,et al.2017.Genetic parameters for growth traits and stem-straightness in Eucalyptus urophylla × E.camaldulensis hybrids from a reciprocal mating design.Euphytica,213(7):142-158.