不同近交程度对湿地松结实与生长的影响
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摘要
【目的】分析近交对广东湿地松遗传材料适应度的影响以及不同亲本的近交衰退差异。【方法】对湿地松C2与C8两个无性系亲本组的16个异交与近交子代开展球果发育与田间生长测定。结实特性主要包括球果形态、种子数量与质量、饱满种子比例等。生长性状主要包括4年生与13年生子代测定林家系树高、胸径和材积。【结果】近交严重降低了两个亲本组单果饱满种子的数量与质量。C8亲本组在F为0.125、0.25和0.5时,饱满种子数量比异交下降了20%、35%和57%,近交衰退ID与近交系数F相关性达0.99。C2亲本组中与F=0.125相比,F为0.25和0.5时饱满种子数下降了50%和67%。保存率在4年生时不表现近交衰退,但13年生保存率总体比异交子代下降了4%(F=0.125)、8%(F=0.25)和19%(F=0.5)。两个亲本组及不同家系对近交的反应不一致,但F=0.125时生长量均表现出严重的近交衰退,13年生时单株材积下降22%。C2组的近交衰退总体大于C8组,13年生林分蓄积量比异交分别下降26%(F=0.125)、20%(F=0.25)和43%(F=0.5)。C8组存在两个生长表现优异的近交家系,表现优于其亲本的自由授粉子代。【结论】湿地松近交子代在球果发育与生长性状上表现近交衰退,在湿地松种子园建设中,应该注重近交管理。不同亲本的近交衰退程度存在差异,湿地松近交子代中也可选育出作为多世代遗传改良的育种材料。
【Objective】Slash pine( Pinus elliottii) was introduced in China 8 decades ago,and has become one of the most important exotic tree species for commercial usage in Guangdong. Inbreeding management is important at multiple cycles of genetic improvement. Most Pinus elliottii breeding programs have progressed to advanced-generation improvement in China. However,limited efforts have been made to evaluate the effects of inbreeding on the important traits of current genetic materials. In current study,the effect of inbreeding on the fitness of Pinus elliottii genetic material in Guangdong and the variation in inbreeding depression( ID) among different parents was analyzed.【Method】C2 and C8 were two super clones chosen based on backward selection. Cone development and field growth were measured in 16 outbreeding and inbred progenies,and in 5 open-pollinated progenies from the C2 and C8 founder clones. There are 4 levels of inbreeding: outcrosses( F = 0),half-sib( F = 0.125),full-sib and backcrosses( F = 0. 25),and selfed( F =0. 5). Seed characteristics including cone length,cone width,seed number per cone,seed weight per cone,filled seeds number per cone,weight of filled seeds per cone,thousand seed weight,proportion of filled seeds and so on,were measured in 1999. The progeny testing stand with a randomized completed block design was established in Taishan,Guangdong in 2000. From 2002,the growth traits were determined,and included height,diameter at breast height ( DBH),individual volume,and plot volume.【Result】Inbreeding hinders cone development in Pinus elliottii,few cones were obtained in 4 inbred families and the rate of seed was obviously reduced at F = 0.5. Furthermore,Inbreeding severely decreased the filled seed number and weight per cone. In C8,the ID at F = 0.125,0.25,and 0.5 was 20%,35%,and 57% for filled seed number,respectively,which was highly related to the inbreeding coefficient( R2= 0.99).In C2,the filled seed number decreased to 50% and 67% at F = 0.25 and 0.5,respectively,when compared to that of F = 0.125. The other traits related to cone morphology and seed production were not consistent in C2 and C8. The ID for survival was not obvious for 4-year-olds,but at F = 0.125,0.25 and 0.5 it was 4%,8% and 19% for survival to 13-year-olds,respectively. There were significant differences between the two founder clones and the related families in growth response to inbreeding. The ID for overall growth was severe at F = 0.125,which was 22% for individual volume at 13 years. The response to inbreeding was higher in C2 than that in C8,and the IDs in C2 followed certain rules: the ID degree was the highest in individual volume,followed by DBH,and the tree height was lowest; The mean IDs in DBH are twice as height. The ID at F = 0.125,0.25,and 0.5 from the C2 founder clone was 13%,15%,and 30% for individual volume at 4 years,respectively,whereas it was 26%,20% and 43% for plot volume at 13 years,respectively. The IDs in C8 were not obvious,and the IDs in plot volume unexpectedly increased by 2.1%. Two specific inbred progenies( C8-2×C8-1 and C8-2×C8-2) showed better growth performance than the C8 and C8-2 open-pollinated progenies.【Conclusion】The inbred progenies exhibited IDs for cone development and growth,which especially caused large economic losses due to the reduction in the survival and plot volume at the mature stage. It was thus indicated that inbreeding management in Pinus elliottii should be given attention during seed orchard establishment. Variations were found in IDs from different parents. The elite inbred progenies could be selected for advanced-generation breeding.
【Objective】Slash pine( Pinus elliottii) was introduced in China 8 decades ago,and has become one of the most important exotic tree species for commercial usage in Guangdong. Inbreeding management is important at multiple cycles of genetic improvement. Most Pinus elliottii breeding programs have progressed to advanced-generation improvement in China. However,limited efforts have been made to evaluate the effects of inbreeding on the important traits of current genetic materials. In current study,the effect of inbreeding on the fitness of Pinus elliottii genetic material in Guangdong and the variation in inbreeding depression( ID) among different parents was analyzed.【Method】C2 and C8 were two super clones chosen based on backward selection. Cone development and field growth were measured in 16 outbreeding and inbred progenies,and in 5 open-pollinated progenies from the C2 and C8 founder clones. There are 4 levels of inbreeding: outcrosses( F = 0),half-sib( F = 0.125),full-sib and backcrosses( F = 0. 25),and selfed( F =0. 5). Seed characteristics including cone length,cone width,seed number per cone,seed weight per cone,filled seeds number per cone,weight of filled seeds per cone,thousand seed weight,proportion of filled seeds and so on,were measured in 1999. The progeny testing stand with a randomized completed block design was established in Taishan,Guangdong in 2000. From 2002,the growth traits were determined,and included height,diameter at breast height ( DBH),individual volume,and plot volume.【Result】Inbreeding hinders cone development in Pinus elliottii,few cones were obtained in 4 inbred families and the rate of seed was obviously reduced at F = 0.5. Furthermore,Inbreeding severely decreased the filled seed number and weight per cone. In C8,the ID at F = 0.125,0.25,and 0.5 was 20%,35%,and 57% for filled seed number,respectively,which was highly related to the inbreeding coefficient( R2= 0.99).In C2,the filled seed number decreased to 50% and 67% at F = 0.25 and 0.5,respectively,when compared to that of F = 0.125. The other traits related to cone morphology and seed production were not consistent in C2 and C8. The ID for survival was not obvious for 4-year-olds,but at F = 0.125,0.25 and 0.5 it was 4%,8% and 19% for survival to 13-year-olds,respectively. There were significant differences between the two founder clones and the related families in growth response to inbreeding. The ID for overall growth was severe at F = 0.125,which was 22% for individual volume at 13 years. The response to inbreeding was higher in C2 than that in C8,and the IDs in C2 followed certain rules: the ID degree was the highest in individual volume,followed by DBH,and the tree height was lowest; The mean IDs in DBH are twice as height. The ID at F = 0.125,0.25,and 0.5 from the C2 founder clone was 13%,15%,and 30% for individual volume at 4 years,respectively,whereas it was 26%,20% and 43% for plot volume at 13 years,respectively. The IDs in C8 were not obvious,and the IDs in plot volume unexpectedly increased by 2.1%. Two specific inbred progenies( C8-2×C8-1 and C8-2×C8-2) showed better growth performance than the C8 and C8-2 open-pollinated progenies.【Conclusion】The inbred progenies exhibited IDs for cone development and growth,which especially caused large economic losses due to the reduction in the survival and plot volume at the mature stage. It was thus indicated that inbreeding management in Pinus elliottii should be given attention during seed orchard establishment. Variations were found in IDs from different parents. The elite inbred progenies could be selected for advanced-generation breeding.
引文
[1]FALCONER D S,MACKAY T F C. Introduction to quantitative genetics[M]. 4th Ed. Essex:Longman Group Limited,1996:65-67.
[2]WU H X,MATHESON A C,SPENCER D. Inbreeding in Pinus radiata. II:time course of inbreeding depression and effect on growth curve[J]. New Zealand Journal of Forestry Science,1998,28(2):123-139.
[3]WU H X,HALLINGBCK H R,SNCHEZ L. Performance of seven tree breeding strategies under conditions of inbreeding depression[J]. G3:Genes,Genomes,Genetics,2016,6(3):529-540.DOI:10.1534/g3.115.025767.
[4]王章荣.高世代种子园营建的一些技术问题[J].南京林业大学学报(自然科学版),2012,36(1):8-10. DOI:10.3969/j.issn.1000-2006,2012.01.002.WANG Z R. Techniques on establishment and management of advanced-generation seed orchard[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2012,36(1):8-10.
[5]FORD G A,MCKEAND S E,JETT J B,et al. Effects of Inbreeding on growth and quality traits in loblolly pine[J]. Forest Science,2015,61(3):579-585.DOI:10.5849/forsci.13-185.
[6]SKROPPA T. Diallel crosses in Picea abies. II:performance and inbreeding depression of selfed line[J]. Forest Genetics,1996,3(2):69-79.
[7]DUREL C E,BERTIN P,KREMER A. Relationship between inbreeding depression and inbreeding coefficient in maritime pine(Pinus pinaster)[J]. Theoretical&Applied Genetics,1996,92(3/4):347-356.DOI:10.1007/bf0023678.
[8] STOEHR M,OTT P,WOODS J. Inbreeding in mid-rotation coastal Douglas-fir:implications for breeding[J]. Annals of Forest Science,2015,72(2):195-204. DOI:10.1007/513595-074-0414-0.
[9]WU H X,MATHESON A C,SPENCER D J,et al. Inbreeding in Pinus radiata. I:the effect of inbreeding on growth,survival and variance[J]. Theoretical and Applied Genetics,1998,97(8):1256-1268.DOI:10.1007/s001220051018.
[10]寇安民,李卓军.油松种子发芽率和苗木高生长的近交效应[J].陕西林业科技,1999(4):31-35.KOU A M,LI Z J. The effect of inbreeding on the germination of seeds and the increment of seedlings of Pinus tabulaeformis[J].Shannxi Forest Science and Technology,1999(4):31-35.
[11]陈益泰,何贵平,李恭学.杉木种子发芽率和苗木高生长的近交效应[J].林业科学研究,1989(5):420-426.CHEN Y T,HE G P,LI G X. The effect of inbreeding on seed germination and seedling height growth of Chinese fir[J]. Forest Research,1989(5):420-426.
[12]潘志刚,游应天.中国主要外来树种引种栽培[M].北京:科学技术出版社,1994:41-42.PAN Z G, YOU Y T. Growing exotic trees in China[M].Beijing:Science and Technology Press,1994:41-42.
[13]雷蕾,潘显强,张露,等.高产脂湿地松松节油成分的遗传变异及综合选择[J].林业科学研究,2015,28(6):804-809.LEI L,PAN X Q,ZHANG L,et al. Genetic variation and comprehensive selection of turpentine composition in high-yielding slash pine(Pinus elliottii)[J]. Forest Research,2015,28(6):804-809.
[14]张耀丽,徐永吉,龙应忠,等.湿地松种植密度对纸浆材主要化学成分的影响[J].南京林业大学学报(自然科学版),2002,26(6):60-62. DOI:10. 3969/j/issn. 1000-2006. 2002.06.016.ZHANG Y L,XU Y J,LONG Y Z,et al. Effects of the content of main chemical composition of slash pine for pulpwood on planting density[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2002,26(6):60-62.
[15]张应中,赵奋成,李福明,等.湿加松F1杂种家系年度生长相关及其早期选择[J].南京林业大学学报(自然科学版),2008,32(4):33-37. DOI:10. 3969/j/issn. 1000-2006. 2008.04.007.ZHANG Y Z,ZHAO F C,LI F M,et al. The growth correlation among ages of Pinus elliottii var. elliottii×P. caribaea var. hondurensis F1 hybrids and its early selection[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(4):33-37.
[16]赵奋成,李宪政,黄永达,等.湿地松改良种子园部分无性系的早期遗传评价[J].广东林业科技,2001,17(2):1-6.ZHAO F C,LI X Z,HUANG Y D,et al. Early genetic analysis of some clones in the improved slash pine seed orchard[J].Guangdong Forestry Science and Technology,2001,17(2):1-6.
[17]MATHESON A C,WHITE T L,POWELL G R,et al. Effects of inbreeding on growth,stem form and rust resistance in Pinus elliottii[J]. Silvae Genetica,1995,44(1):37-46.
[18]朱志淞,李宪政,任海丽,等.台山红岭湿地松种子园的营建和经营管理技术阶段总结[J].广东林业科技,1990(3):1-4.
[19] WEST P W. Tree height[C]//Tree and forest measurement.Berlin,Heidelberg:Springer Berlin Heidelberg,2009:18.
[20]赵奋成,林昌明,吴惠姗,等.湿地松生长性状遗传参数的年度变化趋势[J].广东林业科技,2015,31(5):1-7.ZHAO F C,LIN C M,WU H S,et al. Annual trend in genetic parameters for growth traits of Pinus elliottii[J]. Guangdong Forestry Science and Technology,2015,31(5):1-7.
[21]LITTELL R C,MILLIKEN G A,STROUP W W,et al. SASfor Mixed models[M]. 2nd Ed. Cary NC:SAS Institute Inc,2006:286-297.
[22]樊莉丽,彭方仁,王改萍,等.楸树自交及种内、种间杂交亲和性的细胞学观察[J].南京林业大学学报(自然科学版),2013,37(4):1-7. DOI:10. 3969/j. issn. 1000-2006. 2013.04.001.FAN L L,PENG F R,WANG G P,et al. Cytological observation of fertilization compatibility of Catalpa bungei after self,intraspecific cross and interspecific cross-pollination[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2013,37(4):1-7.
[23]怀特T L,亚当斯W T,尼尔D B.森林遗传学[M].北京:科学出版社,2013:81-85.WHITE T L,ADAMS W T,NEALE D B. Forest genetics[M].Beijing:Science Press,2013:81-85.
[24]SORENSEN F C. Relationship between self-fertility,allocation of growth,and inbreeding depression in three coniferous species[J].Evolution,1999,53(2):417-425.
[25]THEODOROU K,COUVET D. On the expected relationship between inbreeding,fitness,and extinction[J]. Genetics Selection Evolution,2006,38(4):371-387.
[26]HUSBAND B C,SCHEMSKE D W. Evolution of the magnitude and timing of inbreeding depression in plants[J]. Evolution,1996,50(1):54-70. DOI:10.2307/2410780.
[27]WU H X,OWEN J V,ABARQUEZ A,et al. Inbreeding in Pinus radiata. V:the effects of inbreeding on fecundity[J].Silvae Genetica,2004,53(1/2/3/4/5/6):80-87. DOI:10.1515/sg-2004-0015.
[28]BOWER A D,AITKEN S N. Mating system and inbreeding depression in white bark pine(Pinus albicaulis Engelm.)[J]. Tree Genetics&Genomes,2007,3(4):379-388.
[29]PEKKALA N,KNOTT K E,KOTIAHO J S,et al. The effect of inbreeding rate on fitness,inbreeding depression and heterosis over a range of inbreeding coefficients[J]. Evolutionary Applications,2014,7(9):1107-1119. DOI:10.1111/eva.12145.
[30]CHARLESWORTH B,WILLIS J H. The genetics of inbreeding depression[J]. Nature Reviews Genetics,2009,10(11):783-796. DOI:10.1038/nrg2664.
[31]WILLIAMS C G,SAVOLAINEN O. Inbreeding depression in conifers:implications for breeding strategy[J]. Forest Science,1996,42(1):102-117.
[2]WU H X,MATHESON A C,SPENCER D. Inbreeding in Pinus radiata. II:time course of inbreeding depression and effect on growth curve[J]. New Zealand Journal of Forestry Science,1998,28(2):123-139.
[3]WU H X,HALLINGBCK H R,SNCHEZ L. Performance of seven tree breeding strategies under conditions of inbreeding depression[J]. G3:Genes,Genomes,Genetics,2016,6(3):529-540.DOI:10.1534/g3.115.025767.
[4]王章荣.高世代种子园营建的一些技术问题[J].南京林业大学学报(自然科学版),2012,36(1):8-10. DOI:10.3969/j.issn.1000-2006,2012.01.002.WANG Z R. Techniques on establishment and management of advanced-generation seed orchard[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2012,36(1):8-10.
[5]FORD G A,MCKEAND S E,JETT J B,et al. Effects of Inbreeding on growth and quality traits in loblolly pine[J]. Forest Science,2015,61(3):579-585.DOI:10.5849/forsci.13-185.
[6]SKROPPA T. Diallel crosses in Picea abies. II:performance and inbreeding depression of selfed line[J]. Forest Genetics,1996,3(2):69-79.
[7]DUREL C E,BERTIN P,KREMER A. Relationship between inbreeding depression and inbreeding coefficient in maritime pine(Pinus pinaster)[J]. Theoretical&Applied Genetics,1996,92(3/4):347-356.DOI:10.1007/bf0023678.
[8] STOEHR M,OTT P,WOODS J. Inbreeding in mid-rotation coastal Douglas-fir:implications for breeding[J]. Annals of Forest Science,2015,72(2):195-204. DOI:10.1007/513595-074-0414-0.
[9]WU H X,MATHESON A C,SPENCER D J,et al. Inbreeding in Pinus radiata. I:the effect of inbreeding on growth,survival and variance[J]. Theoretical and Applied Genetics,1998,97(8):1256-1268.DOI:10.1007/s001220051018.
[10]寇安民,李卓军.油松种子发芽率和苗木高生长的近交效应[J].陕西林业科技,1999(4):31-35.KOU A M,LI Z J. The effect of inbreeding on the germination of seeds and the increment of seedlings of Pinus tabulaeformis[J].Shannxi Forest Science and Technology,1999(4):31-35.
[11]陈益泰,何贵平,李恭学.杉木种子发芽率和苗木高生长的近交效应[J].林业科学研究,1989(5):420-426.CHEN Y T,HE G P,LI G X. The effect of inbreeding on seed germination and seedling height growth of Chinese fir[J]. Forest Research,1989(5):420-426.
[12]潘志刚,游应天.中国主要外来树种引种栽培[M].北京:科学技术出版社,1994:41-42.PAN Z G, YOU Y T. Growing exotic trees in China[M].Beijing:Science and Technology Press,1994:41-42.
[13]雷蕾,潘显强,张露,等.高产脂湿地松松节油成分的遗传变异及综合选择[J].林业科学研究,2015,28(6):804-809.LEI L,PAN X Q,ZHANG L,et al. Genetic variation and comprehensive selection of turpentine composition in high-yielding slash pine(Pinus elliottii)[J]. Forest Research,2015,28(6):804-809.
[14]张耀丽,徐永吉,龙应忠,等.湿地松种植密度对纸浆材主要化学成分的影响[J].南京林业大学学报(自然科学版),2002,26(6):60-62. DOI:10. 3969/j/issn. 1000-2006. 2002.06.016.ZHANG Y L,XU Y J,LONG Y Z,et al. Effects of the content of main chemical composition of slash pine for pulpwood on planting density[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2002,26(6):60-62.
[15]张应中,赵奋成,李福明,等.湿加松F1杂种家系年度生长相关及其早期选择[J].南京林业大学学报(自然科学版),2008,32(4):33-37. DOI:10. 3969/j/issn. 1000-2006. 2008.04.007.ZHANG Y Z,ZHAO F C,LI F M,et al. The growth correlation among ages of Pinus elliottii var. elliottii×P. caribaea var. hondurensis F1 hybrids and its early selection[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(4):33-37.
[16]赵奋成,李宪政,黄永达,等.湿地松改良种子园部分无性系的早期遗传评价[J].广东林业科技,2001,17(2):1-6.ZHAO F C,LI X Z,HUANG Y D,et al. Early genetic analysis of some clones in the improved slash pine seed orchard[J].Guangdong Forestry Science and Technology,2001,17(2):1-6.
[17]MATHESON A C,WHITE T L,POWELL G R,et al. Effects of inbreeding on growth,stem form and rust resistance in Pinus elliottii[J]. Silvae Genetica,1995,44(1):37-46.
[18]朱志淞,李宪政,任海丽,等.台山红岭湿地松种子园的营建和经营管理技术阶段总结[J].广东林业科技,1990(3):1-4.
[19] WEST P W. Tree height[C]//Tree and forest measurement.Berlin,Heidelberg:Springer Berlin Heidelberg,2009:18.
[20]赵奋成,林昌明,吴惠姗,等.湿地松生长性状遗传参数的年度变化趋势[J].广东林业科技,2015,31(5):1-7.ZHAO F C,LIN C M,WU H S,et al. Annual trend in genetic parameters for growth traits of Pinus elliottii[J]. Guangdong Forestry Science and Technology,2015,31(5):1-7.
[21]LITTELL R C,MILLIKEN G A,STROUP W W,et al. SASfor Mixed models[M]. 2nd Ed. Cary NC:SAS Institute Inc,2006:286-297.
[22]樊莉丽,彭方仁,王改萍,等.楸树自交及种内、种间杂交亲和性的细胞学观察[J].南京林业大学学报(自然科学版),2013,37(4):1-7. DOI:10. 3969/j. issn. 1000-2006. 2013.04.001.FAN L L,PENG F R,WANG G P,et al. Cytological observation of fertilization compatibility of Catalpa bungei after self,intraspecific cross and interspecific cross-pollination[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2013,37(4):1-7.
[23]怀特T L,亚当斯W T,尼尔D B.森林遗传学[M].北京:科学出版社,2013:81-85.WHITE T L,ADAMS W T,NEALE D B. Forest genetics[M].Beijing:Science Press,2013:81-85.
[24]SORENSEN F C. Relationship between self-fertility,allocation of growth,and inbreeding depression in three coniferous species[J].Evolution,1999,53(2):417-425.
[25]THEODOROU K,COUVET D. On the expected relationship between inbreeding,fitness,and extinction[J]. Genetics Selection Evolution,2006,38(4):371-387.
[26]HUSBAND B C,SCHEMSKE D W. Evolution of the magnitude and timing of inbreeding depression in plants[J]. Evolution,1996,50(1):54-70. DOI:10.2307/2410780.
[27]WU H X,OWEN J V,ABARQUEZ A,et al. Inbreeding in Pinus radiata. V:the effects of inbreeding on fecundity[J].Silvae Genetica,2004,53(1/2/3/4/5/6):80-87. DOI:10.1515/sg-2004-0015.
[28]BOWER A D,AITKEN S N. Mating system and inbreeding depression in white bark pine(Pinus albicaulis Engelm.)[J]. Tree Genetics&Genomes,2007,3(4):379-388.
[29]PEKKALA N,KNOTT K E,KOTIAHO J S,et al. The effect of inbreeding rate on fitness,inbreeding depression and heterosis over a range of inbreeding coefficients[J]. Evolutionary Applications,2014,7(9):1107-1119. DOI:10.1111/eva.12145.
[30]CHARLESWORTH B,WILLIS J H. The genetics of inbreeding depression[J]. Nature Reviews Genetics,2009,10(11):783-796. DOI:10.1038/nrg2664.
[31]WILLIAMS C G,SAVOLAINEN O. Inbreeding depression in conifers:implications for breeding strategy[J]. Forest Science,1996,42(1):102-117.