油松不同种源和家系苗期性状的遗传变异分析
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摘要
油松(P.tabulaeformis Carr.)生长快、材质好、耐瘠薄,天然分布于我国北方的14个省(区),是特有的用材树种,也是北方各省区的主要造林树种之一。陕西省具有最丰富的油松遗传资源,全国油松天然林总面积的45%分布在陕西省境内。随着经济发展和国家建设,油松木材的需求量不断增加,但是目前油松优良基因资源损失严重,造林用种数量不足,品质低劣,严重影响了造林效果。在油松天然分布区内开展优良种源,优良家系,优良单株的选择研究,提高油松造林用种的遗传品质具有重要的意义。通过选择优良的基因型,提高目的基因的频率,改变油松群体的结构,增强群体的遗传水平,选育出生长好、产量高、抗性强的优良品系。
本文采集陕西洛南、黄龙、太白、宁陕、黄陵和富县6个种源地的95株油松自由授粉优树种子,采用完全随机区组设计进行容器育苗试验,测定各家系2年生油松实生苗的株高、地径、针叶长、针叶宽、地上部长、主根长、地上部鲜质量、地上部干质量、地下部鲜质量、地下部干质量、侧根数、根冠比、茎干重/茎鲜重、根干重/根鲜重、茎干重/茎长、根干重/根长、可溶性蛋白含量、丙二醛(MDA)和超氧化物歧化酶(SOD),对测定性状进行方差分析、遗传相关分析、遗传变异分析,并采用遗传主成分分析法和综合选择指数法评选优良种源和家系。主要研究结果和结论如下:
1.油松实生苗的各性状,除侧根数在种源间的差异、地茎和主根长在种源间和家系间的差异均不显著外,其他性状在种源间和家系间均达显著或极显著水平。
2.种源间油松实生苗各性状的多重比较表明,洛南种源的株高、针叶长、针叶宽、地上部长、地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量的均值最大,显著高于其他种源的性状的值。黄陵种源的地茎、侧根数和黄龙种源的主根长的值也最大。洛南、太白和富县种源间的可溶性蛋白含量差异达极显著水平,黄龙、太白、宁陕和黄陵种源间的SOD活性差异达显著水平。丙二醛含量在太白和宁陕种源间的差异达极显著水平。
3.油松各家系实生苗在株高、地茎、针叶长、针叶宽、侧根数间遗传相关达到显著或极显著水平,株高与针叶长、针叶宽的表型相关和遗传相关均达极显著水平,相关性一致,而且遗传相关系数均大于表型相关系数。
油松实生苗的地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量4者间的遗传相关系数≥0.8768,油松实生苗地上部长、地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量5者间的表型相关系数均≥0.738,达到极限值水平,同时,以上5性状的遗传相关性和表型相关性一致。
油松实生苗的根冠比与茎干重/茎长、根干重/根长之间的表型相关达极显著负相关,茎干重/茎长与根干重/根长之间的遗传相关均达极显著正相关。SOD与其他性状无显著遗传相关。
4.数量遗传分析表明,可溶性蛋白含量的家系遗传力和单株遗传力均达最大,分别为0.992和0.997,而主根长的家系遗传力和单株遗传力均最小,分别为0.105和0.136,其他性状都受中等或中等偏上遗传力控制。地茎、针叶长、针叶宽、主根长、侧根数、茎干重/茎鲜重和根干重/根鲜重的遗传变异系数≤15.604%,其他性状的遗传变异系数均较大。除主根长和针叶宽外其他性状的遗传增益值均较大,对油松优良家系的选是可行的。
5.经主成分分析,对油松实生苗家系进行综合评价,选择出综合性状表现较好的家系:L33、L49、L44、H34、C1、XY4、H23、T29、Q3、XY(XY4)。根据生长性状和生物量性状综合指数选择出19个优良家系:L47、L20、L29、L26、L2、L8、L14、L49、L50、L25、L33、L44、L15、T23、C1、T11、XY(HY4)、H34和T19。根据抗性性状指数选择出19个优良家系:T23、XY(HY4)、L26、XY(HY3)、H40、L29、XY(6)、H43、L49、L44、H46、L20、XY(HY1)、H38、L33、H34、T29、H15和T31。
Pinus tableaformis Carr. is distributed in the north of China’14 provinces (regions), it is one of the timber tree species and the major species for afforestation ,because of rapid growth、good quality and poor resistance .There is 45% of the total area of natural pine forest in Shaanxi While Shaanxi has the most abundant pine genetic resources. With the economic development and nation-building needs, the demand for pine wood is increasing. However, the situation of loss of genetic resources、lack of tree species and por quality were affected planting effect seriously. we should carry out studies on selection of superior provenance、good family and superior individual , improving the genetic quality of pine. That we have to do lots of works on selecting high quality genotype、increasing the frequency of target gene、changing in forest structure、enhancing the genetic capacity of pine group, to select pine’fine lines with good growth、good quality and strong resistance.
The open pollinated seeds of 95 P. tabulaeformis families collected form Luo Nan、Huang Long、Tai Bai、Ning Shan、Huang Ling and Fu Xian in Shaanxi province; Seedling cultivation was done in container nursery with complete random block design. Determining Height、Ground-diameter、Needle length、Needle width、Shoot length、Main root length、Shoot fresh weight、Shoot dry weight、Root fresh weight、Root dry weight、The number of side root、Root-shoot ratio、Shoot dry weight/shoot fresh weight、Root dry weight/root fresh weight、Shoot dry weight/shoot length、Root dry weight/root length、Soluble protein content、MDA、SOD on 2-years old seedlings and were studied through variance analysis, genetic correlation analysis、genetic variation analysis. And the superior families and individual were selected through principal component analysis and the method of comprehensive selection index. The main results and conclusions were as follows:
1. There were highly significant differences among different families and provenance except ground-diameter and main root length .There was a highly significant difference on the number of side root among families.
2. Multi-comparison of 19 characters among provenances demonstrated that the mean value of height、needle length、needle width、shoot length、shoot fresh weight、shoot dry weight、root fresh weight、root dry weight were the greatest in Luo Nan provenances. The mean value of root-shoot ratio and the number of side root in Huang Ling provenance were the greatest, while the same as the mean value of the main root length in Huang Long provenance. There were highly significant differences on soluble protein content among Luo Nan、Tai Bai and Fu Xian provenances. A significant differences were on SOD among Huang Long、Ning Shan、Tai Bai and Huang Ling provenances.A significant differences were on MDA between Ning Shan and Tai Bai provenances.
3. The coefficients of genetic correlate of seeding of Pinus tabulaeformis were highly significant on height、ground-diameter、needle length、needle width and the number of side root among families . Phenotypic correlation and genetic correlation are all highly significant positive correlation between height and needle length、needle. The coefficients of genetic correlate between height and needle length、needle were greater than the coefficients of phenotypic correlate of that .
The coefficients of genetic correlate about seeding of Pinus tabulaeformis were more than 87.67% among shoot fresh weight, root fresh weight, shoot dry weight and root dry weight. The coefficients of phenotypic correlation were more than 73.8% among root length、shoot fresh weight , root fresh weight, shoot dry weight and root dry weight. Phenotypic correlation and genetic correlation are all highly significant positive correlation among 5 characters above.
Phenotypic correlation were highly negative correlation among root-shoot ratio、Shoot dry weight/shoot length、Root dry weight/root length. Genetic correlation were highly positive correlation between root dry weight/root fresh weight and shoot dry weight/shoot fresh weight , between Shoot dry weight/shoot length and Root dry weight/root length.. There is no significant genetic correlation between SOD and other characters.
4. Quantitative genetic analysis demonstrated that family heritability and single tree heritability of soluble protein content were greater, which were 0.992 and 0.997 respectively. While family heritability and single tree heritability of main root length were lower, which were 0.105 and 0.136 respectively.And other characters were controlled by medium heritability. Genetic variation coefficient of ground-diameter、needle length、needle width、main root length、the number of side root、Shoot dry weight/shoot fresh weight、root dry weight/root fresh weight were less than 15.604%.The characters of seedling Pinus tabulaeformis were greater, except main root length and needle width, which was showed that selecting superior families was a good method to improve the genetic quality of Pinus tabulaeformis.
5. Superior families of seedling Pinus tabuaeformis were evaluated thought principal component analysis. Superior families selected were L33、L49、L44、H34、C1、XY4、H23、T29、Q3、XY(XY4) .According to growth characters and biomass character in index selection. we can determine the index value of and selected 19 superior families, which were L47、L20、L29、L26、L2、L8、L14、L49、L50、L25、L33、L44、L15、T23、C1、T11、XY(HY4)、H34 and T19.According to resistance characters in index selection, we can determine the index value of and selected 19 superior families, which were T23、XY(HY4)、L26、XY(HY3)、H40、L29、XY(6)、H43、L49、L44、H46、L20、XY(HY1)、H38、L33、H34、T29、H15 and T31。
本文采集陕西洛南、黄龙、太白、宁陕、黄陵和富县6个种源地的95株油松自由授粉优树种子,采用完全随机区组设计进行容器育苗试验,测定各家系2年生油松实生苗的株高、地径、针叶长、针叶宽、地上部长、主根长、地上部鲜质量、地上部干质量、地下部鲜质量、地下部干质量、侧根数、根冠比、茎干重/茎鲜重、根干重/根鲜重、茎干重/茎长、根干重/根长、可溶性蛋白含量、丙二醛(MDA)和超氧化物歧化酶(SOD),对测定性状进行方差分析、遗传相关分析、遗传变异分析,并采用遗传主成分分析法和综合选择指数法评选优良种源和家系。主要研究结果和结论如下:
1.油松实生苗的各性状,除侧根数在种源间的差异、地茎和主根长在种源间和家系间的差异均不显著外,其他性状在种源间和家系间均达显著或极显著水平。
2.种源间油松实生苗各性状的多重比较表明,洛南种源的株高、针叶长、针叶宽、地上部长、地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量的均值最大,显著高于其他种源的性状的值。黄陵种源的地茎、侧根数和黄龙种源的主根长的值也最大。洛南、太白和富县种源间的可溶性蛋白含量差异达极显著水平,黄龙、太白、宁陕和黄陵种源间的SOD活性差异达显著水平。丙二醛含量在太白和宁陕种源间的差异达极显著水平。
3.油松各家系实生苗在株高、地茎、针叶长、针叶宽、侧根数间遗传相关达到显著或极显著水平,株高与针叶长、针叶宽的表型相关和遗传相关均达极显著水平,相关性一致,而且遗传相关系数均大于表型相关系数。
油松实生苗的地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量4者间的遗传相关系数≥0.8768,油松实生苗地上部长、地上部鲜质量、地上部干质量、地下部鲜质量和地下部干质量5者间的表型相关系数均≥0.738,达到极限值水平,同时,以上5性状的遗传相关性和表型相关性一致。
油松实生苗的根冠比与茎干重/茎长、根干重/根长之间的表型相关达极显著负相关,茎干重/茎长与根干重/根长之间的遗传相关均达极显著正相关。SOD与其他性状无显著遗传相关。
4.数量遗传分析表明,可溶性蛋白含量的家系遗传力和单株遗传力均达最大,分别为0.992和0.997,而主根长的家系遗传力和单株遗传力均最小,分别为0.105和0.136,其他性状都受中等或中等偏上遗传力控制。地茎、针叶长、针叶宽、主根长、侧根数、茎干重/茎鲜重和根干重/根鲜重的遗传变异系数≤15.604%,其他性状的遗传变异系数均较大。除主根长和针叶宽外其他性状的遗传增益值均较大,对油松优良家系的选是可行的。
5.经主成分分析,对油松实生苗家系进行综合评价,选择出综合性状表现较好的家系:L33、L49、L44、H34、C1、XY4、H23、T29、Q3、XY(XY4)。根据生长性状和生物量性状综合指数选择出19个优良家系:L47、L20、L29、L26、L2、L8、L14、L49、L50、L25、L33、L44、L15、T23、C1、T11、XY(HY4)、H34和T19。根据抗性性状指数选择出19个优良家系:T23、XY(HY4)、L26、XY(HY3)、H40、L29、XY(6)、H43、L49、L44、H46、L20、XY(HY1)、H38、L33、H34、T29、H15和T31。
Pinus tableaformis Carr. is distributed in the north of China’14 provinces (regions), it is one of the timber tree species and the major species for afforestation ,because of rapid growth、good quality and poor resistance .There is 45% of the total area of natural pine forest in Shaanxi While Shaanxi has the most abundant pine genetic resources. With the economic development and nation-building needs, the demand for pine wood is increasing. However, the situation of loss of genetic resources、lack of tree species and por quality were affected planting effect seriously. we should carry out studies on selection of superior provenance、good family and superior individual , improving the genetic quality of pine. That we have to do lots of works on selecting high quality genotype、increasing the frequency of target gene、changing in forest structure、enhancing the genetic capacity of pine group, to select pine’fine lines with good growth、good quality and strong resistance.
The open pollinated seeds of 95 P. tabulaeformis families collected form Luo Nan、Huang Long、Tai Bai、Ning Shan、Huang Ling and Fu Xian in Shaanxi province; Seedling cultivation was done in container nursery with complete random block design. Determining Height、Ground-diameter、Needle length、Needle width、Shoot length、Main root length、Shoot fresh weight、Shoot dry weight、Root fresh weight、Root dry weight、The number of side root、Root-shoot ratio、Shoot dry weight/shoot fresh weight、Root dry weight/root fresh weight、Shoot dry weight/shoot length、Root dry weight/root length、Soluble protein content、MDA、SOD on 2-years old seedlings and were studied through variance analysis, genetic correlation analysis、genetic variation analysis. And the superior families and individual were selected through principal component analysis and the method of comprehensive selection index. The main results and conclusions were as follows:
1. There were highly significant differences among different families and provenance except ground-diameter and main root length .There was a highly significant difference on the number of side root among families.
2. Multi-comparison of 19 characters among provenances demonstrated that the mean value of height、needle length、needle width、shoot length、shoot fresh weight、shoot dry weight、root fresh weight、root dry weight were the greatest in Luo Nan provenances. The mean value of root-shoot ratio and the number of side root in Huang Ling provenance were the greatest, while the same as the mean value of the main root length in Huang Long provenance. There were highly significant differences on soluble protein content among Luo Nan、Tai Bai and Fu Xian provenances. A significant differences were on SOD among Huang Long、Ning Shan、Tai Bai and Huang Ling provenances.A significant differences were on MDA between Ning Shan and Tai Bai provenances.
3. The coefficients of genetic correlate of seeding of Pinus tabulaeformis were highly significant on height、ground-diameter、needle length、needle width and the number of side root among families . Phenotypic correlation and genetic correlation are all highly significant positive correlation between height and needle length、needle. The coefficients of genetic correlate between height and needle length、needle were greater than the coefficients of phenotypic correlate of that .
The coefficients of genetic correlate about seeding of Pinus tabulaeformis were more than 87.67% among shoot fresh weight, root fresh weight, shoot dry weight and root dry weight. The coefficients of phenotypic correlation were more than 73.8% among root length、shoot fresh weight , root fresh weight, shoot dry weight and root dry weight. Phenotypic correlation and genetic correlation are all highly significant positive correlation among 5 characters above.
Phenotypic correlation were highly negative correlation among root-shoot ratio、Shoot dry weight/shoot length、Root dry weight/root length. Genetic correlation were highly positive correlation between root dry weight/root fresh weight and shoot dry weight/shoot fresh weight , between Shoot dry weight/shoot length and Root dry weight/root length.. There is no significant genetic correlation between SOD and other characters.
4. Quantitative genetic analysis demonstrated that family heritability and single tree heritability of soluble protein content were greater, which were 0.992 and 0.997 respectively. While family heritability and single tree heritability of main root length were lower, which were 0.105 and 0.136 respectively.And other characters were controlled by medium heritability. Genetic variation coefficient of ground-diameter、needle length、needle width、main root length、the number of side root、Shoot dry weight/shoot fresh weight、root dry weight/root fresh weight were less than 15.604%.The characters of seedling Pinus tabulaeformis were greater, except main root length and needle width, which was showed that selecting superior families was a good method to improve the genetic quality of Pinus tabulaeformis.
5. Superior families of seedling Pinus tabuaeformis were evaluated thought principal component analysis. Superior families selected were L33、L49、L44、H34、C1、XY4、H23、T29、Q3、XY(XY4) .According to growth characters and biomass character in index selection. we can determine the index value of and selected 19 superior families, which were L47、L20、L29、L26、L2、L8、L14、L49、L50、L25、L33、L44、L15、T23、C1、T11、XY(HY4)、H34 and T19.According to resistance characters in index selection, we can determine the index value of and selected 19 superior families, which were T23、XY(HY4)、L26、XY(HY3)、H40、L29、XY(6)、H43、L49、L44、H46、L20、XY(HY1)、H38、L33、H34、T29、H15 and T31。
引文
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