马尾松二代育种亲本选择及种子园交配系统研究
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摘要
马尾松(Pinus. massoniana)分布广,生长快,耐干旱瘠薄,是我国南方山地主要针叶造林树种之一,广泛用于制浆造纸、建筑用材、采脂等,支撑着我国众多的造纸、木材加工、林产化工等产业发展。本文利用已达1/2轮伐期的巢式交配设计的双亲控制授粉家系遗传测定林,进一步揭示了生长、形质和材性性状所受的遗传控制方式,并为增补二代核心育种亲本筛选了一批速生优质的杂交组合和个体;其次,利用现有二代育种群体及创制的三代育种新种质,揭示了二代育种亲本生长、开花结实特性及与其遗传背景的相关性,探索了三代新种质苗期生长所受遗传控制式样,以为二代种子园建园亲本选择和强杂种优势新种质创制的二代亲本选配提供理论依据和指导;第三,针对国内较早建立且已进入正常开花结实期的二代无性系种子园,研究和揭示了亲本无性系生长、分枝及开花结实的遗传变异规律,了解了二代种子园遗传交配系统及遗传多样性状况,探索了无性系开花结实与种子园交配系统和遗传多样性的相关性,以期为马尾松二代种子园营建和优质高产的遗传经营管理提供重要的科学理论依据。主要研究结果如下:
1、利用12年生的马尾松巢式交配设计的遗传测定林材料研究表明,除树干通直度外,不同杂交组合间、父本间以及相同父本不同母本间的生长与木材基本密度等存在显著的遗传差异。胸径、树高和单株材积及树干圆满度同时受到父本和母本效应的影响,且两者效应差异较小,而木材基本密度则主要受母本效应的影响,其母本效应是父本效应的1.56倍。马尾松胸径和木材基本密度以加性效应控制为主,显性效应次之,而树高、单株材积和树干圆满度几乎完全由加性效应控制,可依据亲本表现对其子代进行预测。比较分析发现,不同杂交组合和父本无性系其生长与木材基本密度的相关性较小,速生的杂交组合和父本无性系并不一定具有较低木材基本密度。各杂交组合生长表现与父本无性系GCA之间相关性不明显,但选配GCA高的父本无性系将会提高获得优良组合的机率。以干物质积累量和胸径等为选择指标,分别选出纸浆纤维材和锯材各10个优良杂交组合,及17个和13个优良单株(即二代育种亲本)。
2、51个马尾松二代育种亲本无性系,7年树龄平均树高、胸径和材积分别为5.53 m,9.05 cm和0.021 m~3,马尾松二代育种亲本生长性状总体上优良,但生长性状在亲本无性系之间存在显著差异,表明二代育种群体具有丰富的遗传变异。对进入开花结实盛期的39个无性系调查发现,开花、结实等生殖性状在不同亲本无性系之间的变异程度高于生长性状。基于主要生长和生殖性状,将39个亲本无性系聚在4个类群,不同类群存在显著性状差异。类群1生长性状总体较优,雌球花、球果量也较多。类群2生长性状总体较差,开花、球果量也较少。类群3生长性状较优、雄球花数量较多,但雌球花、球果数量较少。类群4生长性状总体较差,但雌球花、球果数多。无性系的雌球花、球果数与生长量呈弱度负相关。39个亲本无性系中,共选出28个生长、生殖性状较为均衡(生长量、开花结实量均在中等或中等以上)的无性系,可将这些无性系作为候选优良亲本,结合亲本配合力的测评,从中选择优良的杂交亲本及二代种子园的建园材料。
3、15个ISSR标记在36个二代亲本无性系中检测到,Nei’s基因多样度(h)为0.328,Shannon信息多样性指数(I)为0.476,亲本间分子遗传距离的范围是0.24～0.82,平均为0.414,在分子水平上证明马尾松二代育种群体遗传多样性较高,具有较好的高世代遗传改良潜力。基于Nei’s无偏遗传距离进行UPGMA聚类,36个无性系中的35个聚在5个类群内,聚类结果与马尾松二代亲本的遗传背景、系谱关系及产地纬度部分吻合。综合认为,在兼顾亲本配合力测定和遗传背景的基础上,优先选配来自不同聚类类群且在一定遗传距离范围内(0.264～0.529)的分子遗传距离较大的杂交亲本,将更利于提高子代的生长表现和杂种优势程度。
4、基于二代育种亲本创制的6×6半双列交配设计三代育种新种质的配合力分析表明,除苗高线性生长期(LGD)外,苗高、地径、苗高最大生长速率(MGR)、线性生长量(TLG)和线性生长速率(LGR)的GCA和SCA均差异显著或极显著,苗高、MGR、LGD和LGR以显性基因效应控制为主,加性基因效应次之,地径几乎完全由显性基因效应控制,而TLG受加性基因效应控制略大,可能与试验亲本经过GCA间接选择有关。与亲本经GCA初步测定的二代种质研究结果相比,马尾松三代种质苗木生长性状所受遗传控制式样变化较小,只是其SCA效应有了明显提高。在马尾松二代亲本选配中,应加强亲本间特殊配合能力的筛选,以创制优良的杂交组合。
5、对于截干矮化马尾松二代无性系种子园,其雌、雄球花量及球果数在无性系内分株间差异不显著,但在无性系间均达极显著水平,其遗传变异系数变化在37.85%～53.50%,无性系重复力皆在0.71以上。根据开花结实的调查结果,种子园无性系主要聚为雌多雄少(偏雌)、雄多雌少(偏雄)、雌雄均多(偏中)和雌雄均少(偏少)4种类型,其分别占总无性系的25.00%、29.55%、38.64%和6.82%,种子园中20%的高产无性系提供了开花结实总量的41.27%～49.15%。相关分析结果认为,理想的截干矮化马尾松二代高产无性系母树的主要形态特征为:冠幅宽、分枝长、分枝角大、树冠宽卵圆形或圆球形。在种实性状上,除种子千粒重外,球果大小和出籽性状在母树树冠阴面与阳面间差异极显著,且均以树冠阳面高于阴面,说明充足的光照有利于提高种子园种子产量。
6、12对SSR引物在马尾松二代无性系种子园中检测到,亲本和子代群体的总平均观测杂合度(Ho)、期望杂合度(He)及多态信息含量(PIC)分别为0.402、0.446和0.399。平均每个采种母树与22个父本产生子代。在自由授粉状态下,马尾松二代种子园自交率为1.56%,自交现象很弱,其交配方式以异交为主。绝大多数亲本的雄性繁殖适合度在1.00%～3.00%之间,平均为2.00%,形成6.4个后代。马尾松有效花粉的散布距离符合正态分布,其主要散布距离集中在0～90 m,而检测到的最大散布距离为185 m。种子园花粉污染率较低,仅为4.06%。总体上,树冠南面子代亲本交配距离较北面有增加的趋势,但树冠南、北面子代父本组成数并未表现某一明显的规律。
7、马尾松二代种子园子代群体包含了亲本群体的所有等位基因,子代与亲本具有同样高的遗传多样性,子代群体的F为0.046,纯合子过剩的现象不明显;树冠北面子代遗传多样性并未因其雌、雄球花量较树冠南面的减少而有明显的降低;雌雄均衡型和偏雌型植株子代遗传多样性基本相当,以偏雌型植株子代略大,两者F均趋于零,基本符合哈温平衡;种子园多位点异交率为1.097,异交率较高,亲本的近交现象不显著(tm-ts=-0.031);树冠南面多位点与单位点异交率均高于树冠北面;多位点异交率在偏雌与雌雄均衡型植株中基本相当,雌雄均衡型植株并未因其雌、雄球花量比例较偏雌型植株减小而呈现异交率明显降低的现象,两种类型植株的近交指数均接近于零。整体而言,研究的马尾松二代种子园生产的良种在遗传多样性上并没有因亲本受人工选择强度的增加而降低,种子园在营建时家系选择和配置均比较成功,无性系间基因交流相对充分,亲本近交现象不明显。
Masson pine (P. massoniana) is one of the conifers native tree species in south china with better quality of fast growth, wide distribution and strong resistance, and used widely in papermaking, building and rosin. In this thesis, 12 years old (1/2 rotation) of P. massoniana with nested mating design was studied to reveal the mode of genetic control in growth and wood quality, some superior crossing combinations and superior individuals were screened for adding to second-generation breeding population of masson pine. The genetic variation of major growth and reproductive traits and the correlation with the genetic background of breeding parents were researched by breeding population and third-generation germplasm formulated by second-generation breeding parents, and the mode of genetic control in seedling growth were also investigated, which would provide theoretical basis and guidance for the selection of seed orchard parents and the formulation of excellent new germplasm based on the matching for second-generation parents. In addition, the genetic variation law of major growth, branching and reproductive traits was investigated in a dwarfed second generation clonal seed orchard of masson pine which had entered into normal flowering and seed setting stage, and the genetic mating system, genetic diversity and the correlation with flowering and fruiting traits were also analysised to provide important theories for genetic management in higher quality and yield of the second generation seed orchard of masson pine .The results are as follows:
1. Male and female parents effects, additive effect and dominant effect of growth traits and wood basic density were investigated in a progeny test forestry of 12 years old of P. massoniana with nested mating design.The results showed that, except for stem straightness, there existed significant differences in growth rate and wood quality among the crossing combinations, male and female/male parents. Male and female parents effects worked on DBH, height, individual volume and stem fullness simultaneously, but the difference of effects was very little. For wood basic density, female parent effect was 1.56 times larger than male parents. DBH and wood basic density of P. massoniana were controlled by additive effect primarily, and then dominant effect, but height, individual volume and stem fullness were almost controlled by additive effect completely, progeny could be predicted by the performance of parents.The result of comparative analysis for growth and wood quality of crossing combinations and male parents clones showed that the correlation between growth and wood basic density was very little, for fast-growing male parents and crossing combinations, the wood basic density was not always low. The correlation between growth performance of crossing combinations and GCA of male parents was not obvious, but the probability to get superior crossing combinations would be raised through selecting male parent with high GCA to hybridizing. Ten superior crossing combinations and 17 and 13 superior individuals for pulp wood and saw timber were screened based on wood dry matter and DBH respectively.
2. The average height, DBH and volume of tested 51 clones in the second generation breeding population were 5.53 m, 9.05 cm and 0.021 m3, respectively, the overall growth performance of masson pine was superior, and significant deviation in growth traits was detected between clones, phenotype genetic variation was abundant in the second generation breeding population. For reproductive traits such as cones number, the degree of variation was higher than growth traits such as height and DBH. Base on major growth traits and reproductive traits, the 39 tested clones were clustered into 4 groups, which exhibited large degree of variation in traits. Group 1 exhibited superior growth traits and large amount of female cones and young cones, whereas group 2 exhibited poor growth traits and cones amounts. In group 3, the growth traits and male cone amount was superior, but the female cone amount was poor. Group 4 exhibited poor growth traits but large amount of female cones and young cones. There was a weak negative correlation between growth traits and female cone amounts. From 39 tested clones, 28 superior clones were selected, which could be used for controlled pollination and seed orchard parents when combined the result of combining ability.
3. 36 parental clones from the second generation breeding population of masson pine were assessed to detect their genetic variation at molecular level using 15 selected ISSR (inter-simple sequence repeat) markers, the Nei's genetic diversity was 0.328, the Shannon's information diversity index was 0.476, and the genetic distances ranged from 0.24～0.82 with an average genetic distance of 0.414, indicated that the genetic variation of the second generation breeding population was at high level, and had the potency for advanced-generation genetic improvement. The UPGMA clustering analysis based on genetic distance were done, the 36 clones was clustered into 5 groups, and the clustering was partially consistent with the background and genealogy of the clones. It was helpful for improving the growth performance and heterosis level of progenies to select cross parents from different groups and some genetic distance range (0.264～0.529) which had greater molecular genetic distance on the basis of the combining ability and background.
4. Combining ability of seedling growth and shoot elongation parameters for the third generation germplasm of P. massoniana were investigated in a 6×6 half diallel cross design, which the parents were selected from the second generation breeding population. The result showed that, except for linear growth day(LGD), there existed significant differences in general combining ability (GCA) and specific combining ability (SCA) of seedling height, ground diameter, maximum growth rate (MGR), linear growth rate (LGR) and total linear growth (TLG), and the dominant gene effect was predominant over the additive gene effect for seedling height, MGR, LGR and LGD, especially for ground diameter, which was almost controlled by dominant gene effect completely, but for TLG, the additive gene effect was somewhat larger than dominant gene effect. The reason for this seemed to be that the indirect selection on GCA of growth had been made for mating parents.Compared to the second generation germplasm, which the mating parents had been indirect selected on GCA of growth, the change was little of genetic control model for this traits in the third generation germplasm, only the variance component of SCA had increased significantly. During the genetic improvement of the advanced generation in P. massoniana, the selection of SCA between parents should be strengthened, which would help to produce superior cross combinations.
5. The genetic variation law of flowering and fruiting was investigated in the dwarfed second generation clonal seed orchard of P. massoniana which had entered into normal flowering and seed setting stage. The results showed that there existed significant differences in strobilus and cone production among the different clones, but there were not among clonal ramets, the genetic variation coefficient varied from 37.85% to 53.50%, and the clone repeatability was over 0.71. The clones could be clustered into four types, partial female type, partial male type, neutral type and few type according to the law of flowering and fruiting, which accounted for 25.00%, 29.55%, 38.64% and 6.82% respectively. About 20% high yield clones provided strobilus and cone for the whole seed orchard production from 41.27% to 49.15%. The results of correlation analysis showed that the optimum mother tree form traits of the dwarfed second generation seed orchard clones with high yield were as follows: larger crown, longer branch, flatter branch angle and wide egg-shaped crown or spheroidal crown etc.. Except for thousand-seed weight, there were significant differences between shaded halves and exposed halves of tree crown in the traits of cone and seed yield, and exposed halves was superior to shaded halves in the traits, which showed that ample illumination would help to improve the yield of seed orchard.
6. The patterns of effective pollen dispersal were explored in an experimental population of the second generation clonal seed orchard of P. massoniana. The average observed and expected heterozygosity (Ho and He) are 0.402 and 0.446 respectively in experimental population. The average polymorphic information content (PIC) is 0.399. Every mother tree with 22 male parents produced progenies. The self-crossing rate reached 1.56% in open-pollination condition, and outcrossing was primary mating mode of the second generation clonal seed orchard of P. massoniana. The reproduction contribution was changed from 1% to 3% for the most of male parents, with average of 2.00% (produced 6.4 progenies). The most pollination events were detected in the seed orchard of P. massoniana ranged from 0 to 90 m, the maximum dispersal distance reached 185 m, and accord with normal distribution. The pollen contamination was only 4.06%. For the mating distance among parents, there was an increasing tend in the south progenies of crown compared with the north, but no obvious trend arised in the male parent numbers of the south and north progenies of crown.
7. The level of genetic diversity in progeny population was the same as in maternal population in the second generation clonal seed orchard of P. massoniana, all alleles detected in progeny population were included in maternal population, there were not obvious surplus phenomenon of homozygotes in progeny population (F=0.047). The genetic diversity was not decreasing tend with reducing of strobilus in the north crown compared with the south crown. The progeny of neutral tree had the same level of genetic diversity as in the progeny of partial female tree, which the fixation index tended to be zero and accorded with Hardy-Weinberg equilibrium. The outcrossing rate of the seed orchard was very high, with 1.097, and biparental inbreeding was not significant (tm-ts=-0.031). The multilocus outcrossing rate and singlelocus outcrossing rate in the south crown were higher than the north crown. The multilocus outcrossing rate in progeny of partial female tree was the same as in progeny of neutral tree, and the outcrossing rate was not obvious decreasing tend with proportional reducing of female and male strobilus in neutral tree compared with partial female tree, which the fixation index tended to be zero. The level of genetic diversity and outcrossing rate in the second clonal seed orchard did not decrease with the increasing of artificial selection strength, gene exchange among clones was sufficient relatively, and biparental inbreeding was not significant.
1、利用12年生的马尾松巢式交配设计的遗传测定林材料研究表明,除树干通直度外,不同杂交组合间、父本间以及相同父本不同母本间的生长与木材基本密度等存在显著的遗传差异。胸径、树高和单株材积及树干圆满度同时受到父本和母本效应的影响,且两者效应差异较小,而木材基本密度则主要受母本效应的影响,其母本效应是父本效应的1.56倍。马尾松胸径和木材基本密度以加性效应控制为主,显性效应次之,而树高、单株材积和树干圆满度几乎完全由加性效应控制,可依据亲本表现对其子代进行预测。比较分析发现,不同杂交组合和父本无性系其生长与木材基本密度的相关性较小,速生的杂交组合和父本无性系并不一定具有较低木材基本密度。各杂交组合生长表现与父本无性系GCA之间相关性不明显,但选配GCA高的父本无性系将会提高获得优良组合的机率。以干物质积累量和胸径等为选择指标,分别选出纸浆纤维材和锯材各10个优良杂交组合,及17个和13个优良单株(即二代育种亲本)。
2、51个马尾松二代育种亲本无性系,7年树龄平均树高、胸径和材积分别为5.53 m,9.05 cm和0.021 m~3,马尾松二代育种亲本生长性状总体上优良,但生长性状在亲本无性系之间存在显著差异,表明二代育种群体具有丰富的遗传变异。对进入开花结实盛期的39个无性系调查发现,开花、结实等生殖性状在不同亲本无性系之间的变异程度高于生长性状。基于主要生长和生殖性状,将39个亲本无性系聚在4个类群,不同类群存在显著性状差异。类群1生长性状总体较优,雌球花、球果量也较多。类群2生长性状总体较差,开花、球果量也较少。类群3生长性状较优、雄球花数量较多,但雌球花、球果数量较少。类群4生长性状总体较差,但雌球花、球果数多。无性系的雌球花、球果数与生长量呈弱度负相关。39个亲本无性系中,共选出28个生长、生殖性状较为均衡(生长量、开花结实量均在中等或中等以上)的无性系,可将这些无性系作为候选优良亲本,结合亲本配合力的测评,从中选择优良的杂交亲本及二代种子园的建园材料。
3、15个ISSR标记在36个二代亲本无性系中检测到,Nei’s基因多样度(h)为0.328,Shannon信息多样性指数(I)为0.476,亲本间分子遗传距离的范围是0.24～0.82,平均为0.414,在分子水平上证明马尾松二代育种群体遗传多样性较高,具有较好的高世代遗传改良潜力。基于Nei’s无偏遗传距离进行UPGMA聚类,36个无性系中的35个聚在5个类群内,聚类结果与马尾松二代亲本的遗传背景、系谱关系及产地纬度部分吻合。综合认为,在兼顾亲本配合力测定和遗传背景的基础上,优先选配来自不同聚类类群且在一定遗传距离范围内(0.264～0.529)的分子遗传距离较大的杂交亲本,将更利于提高子代的生长表现和杂种优势程度。
4、基于二代育种亲本创制的6×6半双列交配设计三代育种新种质的配合力分析表明,除苗高线性生长期(LGD)外,苗高、地径、苗高最大生长速率(MGR)、线性生长量(TLG)和线性生长速率(LGR)的GCA和SCA均差异显著或极显著,苗高、MGR、LGD和LGR以显性基因效应控制为主,加性基因效应次之,地径几乎完全由显性基因效应控制,而TLG受加性基因效应控制略大,可能与试验亲本经过GCA间接选择有关。与亲本经GCA初步测定的二代种质研究结果相比,马尾松三代种质苗木生长性状所受遗传控制式样变化较小,只是其SCA效应有了明显提高。在马尾松二代亲本选配中,应加强亲本间特殊配合能力的筛选,以创制优良的杂交组合。
5、对于截干矮化马尾松二代无性系种子园,其雌、雄球花量及球果数在无性系内分株间差异不显著,但在无性系间均达极显著水平,其遗传变异系数变化在37.85%～53.50%,无性系重复力皆在0.71以上。根据开花结实的调查结果,种子园无性系主要聚为雌多雄少(偏雌)、雄多雌少(偏雄)、雌雄均多(偏中)和雌雄均少(偏少)4种类型,其分别占总无性系的25.00%、29.55%、38.64%和6.82%,种子园中20%的高产无性系提供了开花结实总量的41.27%～49.15%。相关分析结果认为,理想的截干矮化马尾松二代高产无性系母树的主要形态特征为:冠幅宽、分枝长、分枝角大、树冠宽卵圆形或圆球形。在种实性状上,除种子千粒重外,球果大小和出籽性状在母树树冠阴面与阳面间差异极显著,且均以树冠阳面高于阴面,说明充足的光照有利于提高种子园种子产量。
6、12对SSR引物在马尾松二代无性系种子园中检测到,亲本和子代群体的总平均观测杂合度(Ho)、期望杂合度(He)及多态信息含量(PIC)分别为0.402、0.446和0.399。平均每个采种母树与22个父本产生子代。在自由授粉状态下,马尾松二代种子园自交率为1.56%,自交现象很弱,其交配方式以异交为主。绝大多数亲本的雄性繁殖适合度在1.00%～3.00%之间,平均为2.00%,形成6.4个后代。马尾松有效花粉的散布距离符合正态分布,其主要散布距离集中在0～90 m,而检测到的最大散布距离为185 m。种子园花粉污染率较低,仅为4.06%。总体上,树冠南面子代亲本交配距离较北面有增加的趋势,但树冠南、北面子代父本组成数并未表现某一明显的规律。
7、马尾松二代种子园子代群体包含了亲本群体的所有等位基因,子代与亲本具有同样高的遗传多样性,子代群体的F为0.046,纯合子过剩的现象不明显;树冠北面子代遗传多样性并未因其雌、雄球花量较树冠南面的减少而有明显的降低;雌雄均衡型和偏雌型植株子代遗传多样性基本相当,以偏雌型植株子代略大,两者F均趋于零,基本符合哈温平衡;种子园多位点异交率为1.097,异交率较高,亲本的近交现象不显著(tm-ts=-0.031);树冠南面多位点与单位点异交率均高于树冠北面;多位点异交率在偏雌与雌雄均衡型植株中基本相当,雌雄均衡型植株并未因其雌、雄球花量比例较偏雌型植株减小而呈现异交率明显降低的现象,两种类型植株的近交指数均接近于零。整体而言,研究的马尾松二代种子园生产的良种在遗传多样性上并没有因亲本受人工选择强度的增加而降低,种子园在营建时家系选择和配置均比较成功,无性系间基因交流相对充分,亲本近交现象不明显。
Masson pine (P. massoniana) is one of the conifers native tree species in south china with better quality of fast growth, wide distribution and strong resistance, and used widely in papermaking, building and rosin. In this thesis, 12 years old (1/2 rotation) of P. massoniana with nested mating design was studied to reveal the mode of genetic control in growth and wood quality, some superior crossing combinations and superior individuals were screened for adding to second-generation breeding population of masson pine. The genetic variation of major growth and reproductive traits and the correlation with the genetic background of breeding parents were researched by breeding population and third-generation germplasm formulated by second-generation breeding parents, and the mode of genetic control in seedling growth were also investigated, which would provide theoretical basis and guidance for the selection of seed orchard parents and the formulation of excellent new germplasm based on the matching for second-generation parents. In addition, the genetic variation law of major growth, branching and reproductive traits was investigated in a dwarfed second generation clonal seed orchard of masson pine which had entered into normal flowering and seed setting stage, and the genetic mating system, genetic diversity and the correlation with flowering and fruiting traits were also analysised to provide important theories for genetic management in higher quality and yield of the second generation seed orchard of masson pine .The results are as follows:
1. Male and female parents effects, additive effect and dominant effect of growth traits and wood basic density were investigated in a progeny test forestry of 12 years old of P. massoniana with nested mating design.The results showed that, except for stem straightness, there existed significant differences in growth rate and wood quality among the crossing combinations, male and female/male parents. Male and female parents effects worked on DBH, height, individual volume and stem fullness simultaneously, but the difference of effects was very little. For wood basic density, female parent effect was 1.56 times larger than male parents. DBH and wood basic density of P. massoniana were controlled by additive effect primarily, and then dominant effect, but height, individual volume and stem fullness were almost controlled by additive effect completely, progeny could be predicted by the performance of parents.The result of comparative analysis for growth and wood quality of crossing combinations and male parents clones showed that the correlation between growth and wood basic density was very little, for fast-growing male parents and crossing combinations, the wood basic density was not always low. The correlation between growth performance of crossing combinations and GCA of male parents was not obvious, but the probability to get superior crossing combinations would be raised through selecting male parent with high GCA to hybridizing. Ten superior crossing combinations and 17 and 13 superior individuals for pulp wood and saw timber were screened based on wood dry matter and DBH respectively.
2. The average height, DBH and volume of tested 51 clones in the second generation breeding population were 5.53 m, 9.05 cm and 0.021 m3, respectively, the overall growth performance of masson pine was superior, and significant deviation in growth traits was detected between clones, phenotype genetic variation was abundant in the second generation breeding population. For reproductive traits such as cones number, the degree of variation was higher than growth traits such as height and DBH. Base on major growth traits and reproductive traits, the 39 tested clones were clustered into 4 groups, which exhibited large degree of variation in traits. Group 1 exhibited superior growth traits and large amount of female cones and young cones, whereas group 2 exhibited poor growth traits and cones amounts. In group 3, the growth traits and male cone amount was superior, but the female cone amount was poor. Group 4 exhibited poor growth traits but large amount of female cones and young cones. There was a weak negative correlation between growth traits and female cone amounts. From 39 tested clones, 28 superior clones were selected, which could be used for controlled pollination and seed orchard parents when combined the result of combining ability.
3. 36 parental clones from the second generation breeding population of masson pine were assessed to detect their genetic variation at molecular level using 15 selected ISSR (inter-simple sequence repeat) markers, the Nei's genetic diversity was 0.328, the Shannon's information diversity index was 0.476, and the genetic distances ranged from 0.24～0.82 with an average genetic distance of 0.414, indicated that the genetic variation of the second generation breeding population was at high level, and had the potency for advanced-generation genetic improvement. The UPGMA clustering analysis based on genetic distance were done, the 36 clones was clustered into 5 groups, and the clustering was partially consistent with the background and genealogy of the clones. It was helpful for improving the growth performance and heterosis level of progenies to select cross parents from different groups and some genetic distance range (0.264～0.529) which had greater molecular genetic distance on the basis of the combining ability and background.
4. Combining ability of seedling growth and shoot elongation parameters for the third generation germplasm of P. massoniana were investigated in a 6×6 half diallel cross design, which the parents were selected from the second generation breeding population. The result showed that, except for linear growth day(LGD), there existed significant differences in general combining ability (GCA) and specific combining ability (SCA) of seedling height, ground diameter, maximum growth rate (MGR), linear growth rate (LGR) and total linear growth (TLG), and the dominant gene effect was predominant over the additive gene effect for seedling height, MGR, LGR and LGD, especially for ground diameter, which was almost controlled by dominant gene effect completely, but for TLG, the additive gene effect was somewhat larger than dominant gene effect. The reason for this seemed to be that the indirect selection on GCA of growth had been made for mating parents.Compared to the second generation germplasm, which the mating parents had been indirect selected on GCA of growth, the change was little of genetic control model for this traits in the third generation germplasm, only the variance component of SCA had increased significantly. During the genetic improvement of the advanced generation in P. massoniana, the selection of SCA between parents should be strengthened, which would help to produce superior cross combinations.
5. The genetic variation law of flowering and fruiting was investigated in the dwarfed second generation clonal seed orchard of P. massoniana which had entered into normal flowering and seed setting stage. The results showed that there existed significant differences in strobilus and cone production among the different clones, but there were not among clonal ramets, the genetic variation coefficient varied from 37.85% to 53.50%, and the clone repeatability was over 0.71. The clones could be clustered into four types, partial female type, partial male type, neutral type and few type according to the law of flowering and fruiting, which accounted for 25.00%, 29.55%, 38.64% and 6.82% respectively. About 20% high yield clones provided strobilus and cone for the whole seed orchard production from 41.27% to 49.15%. The results of correlation analysis showed that the optimum mother tree form traits of the dwarfed second generation seed orchard clones with high yield were as follows: larger crown, longer branch, flatter branch angle and wide egg-shaped crown or spheroidal crown etc.. Except for thousand-seed weight, there were significant differences between shaded halves and exposed halves of tree crown in the traits of cone and seed yield, and exposed halves was superior to shaded halves in the traits, which showed that ample illumination would help to improve the yield of seed orchard.
6. The patterns of effective pollen dispersal were explored in an experimental population of the second generation clonal seed orchard of P. massoniana. The average observed and expected heterozygosity (Ho and He) are 0.402 and 0.446 respectively in experimental population. The average polymorphic information content (PIC) is 0.399. Every mother tree with 22 male parents produced progenies. The self-crossing rate reached 1.56% in open-pollination condition, and outcrossing was primary mating mode of the second generation clonal seed orchard of P. massoniana. The reproduction contribution was changed from 1% to 3% for the most of male parents, with average of 2.00% (produced 6.4 progenies). The most pollination events were detected in the seed orchard of P. massoniana ranged from 0 to 90 m, the maximum dispersal distance reached 185 m, and accord with normal distribution. The pollen contamination was only 4.06%. For the mating distance among parents, there was an increasing tend in the south progenies of crown compared with the north, but no obvious trend arised in the male parent numbers of the south and north progenies of crown.
7. The level of genetic diversity in progeny population was the same as in maternal population in the second generation clonal seed orchard of P. massoniana, all alleles detected in progeny population were included in maternal population, there were not obvious surplus phenomenon of homozygotes in progeny population (F=0.047). The genetic diversity was not decreasing tend with reducing of strobilus in the north crown compared with the south crown. The progeny of neutral tree had the same level of genetic diversity as in the progeny of partial female tree, which the fixation index tended to be zero and accorded with Hardy-Weinberg equilibrium. The outcrossing rate of the seed orchard was very high, with 1.097, and biparental inbreeding was not significant (tm-ts=-0.031). The multilocus outcrossing rate and singlelocus outcrossing rate in the south crown were higher than the north crown. The multilocus outcrossing rate in progeny of partial female tree was the same as in progeny of neutral tree, and the outcrossing rate was not obvious decreasing tend with proportional reducing of female and male strobilus in neutral tree compared with partial female tree, which the fixation index tended to be zero. The level of genetic diversity and outcrossing rate in the second clonal seed orchard did not decrease with the increasing of artificial selection strength, gene exchange among clones was sufficient relatively, and biparental inbreeding was not significant.
引文
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