长白落叶松优树子代生长与材质的遗传变异及多性状联合选择
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摘要
本文以黑龙江省林业科学院江山娇实验林场24年生长白落叶松优树子代测定林为研究对象,采用胸径木芯与伐倒木2种取样方法,对14个处理的生长性状、木芯材质性状(4个性状)、解析木材质性状(21个性状)和制浆造纸特性(16个性状)以及木材物理力学性状(20个性状)进行了遗传变异分析、方差分析、相关分析、综合指数选择研究。摸索出家系间生长性状、木材材性性状的遗传变异规律,研究了性状间的相关关系,建立了木芯基本密度、管胞性状与解析木相应性状的回归模型,利用综合指数法选出了2个纸浆材和建筑材兼用的优良家系。为初级无性系种子园疏伐与升级改建提供理论依据,为杂交育种亲本选择以及高世代种子园优树选择打下了良好基础。主要研究结果如下：
1.性状的遗传变异：生长性状、木芯基本密度和管胞性状、解析木材质性状和制浆造纸性状以及木材物理力学强度均存在较大的变异,家系内个体间也存在着丰富的变异。家系间生长性状差异极显著,树高、胸径和材积的家系遗传力分别为0.73、0.72和0.80。家系间木芯基本密度、木芯管胞性状差异显著,家系遗传力在0.56-0.80之间。解析木早材微纤丝角、管胞长度、早材壁腔比、木质素、阿拉伯糖含量、抗张强度、撕裂度等家系间差异极显著,家系遗传力在0.63-0.86之间,基本密度、晚材率、管胞长宽比、综纤维素含量等家系间差异显著,家系遗传力在0.51-0.61；家系间弦面抗劈力、端面硬度和弦面硬度差异极显著,家系遗传力分别为0.77、0.62和0.72；家系间气干密度、晚材率、气干体积干缩率、径面抗劈力、抗弯弹性模量和径面顺纹抗剪强度差异显著,家系遗传力在0.52-0.60之间,这些性状受中等及以上强度的遗传控制,家系水平的材性改良潜力较大
2.性状相关分析：牛长性状与管胞长度呈极显著正相关,且相关关系主要受遗传机制控制,与木材密度、管胞长宽比相关不显著,与大多数木材力学强度指标相关不显著,可以进行独立选择；木材密度与晚材率、径向干缩率、弦面抗劈力、抗弯弹性模量、抗弯强度、径面顺纹抗剪强度、顺纹抗压强度、硬度(端面、径面)呈极显著或显著正相关,与纸浆材其它性状相关不显著,说明木材密度的改良有利于木材物理力学性状的综合改良,与纸浆材性状可以独立选择；管胞长宽比与早材弦向直径、纸的抗张强度、抗张指数、撕裂度和撕裂指数呈极显著或显著正相关,与管胞比量呈极显著负相关。综纤维素含量与粗浆得率、细浆得率呈极显著正相关,与苯醇抽出物含量、冷水抽提物含量、阿拉伯糖含量和浆的硬度呈极显著或显著负相关。早材纤丝角与早材双壁厚呈极显著正相关,与管胞长度和管胞长宽比、撕裂度和撕裂指数、木材抗弯弹性模量呈显著负相关；晚材率与抗劈力、抗弯弹性模量、抗弯强度、顺纹抗压强度呈极显著正相关,与早材壁腔比、浆料纤维长宽比、木材端而硬度呈显著正相关,与早材径向直径、树脂道比量、浆料纤维宽度、浆料壁腔比、抗张强度、抗张指数、耐破度和耐破指数呈极显著或显著负相关。
3.木芯基本密度、管胞长度、管胞宽度以及管胞长宽比与解析木相应性状呈极显著正相关关系,且回归模型较为理想,可以利用胸径木芯各性状值预测其单株值,间接选择和评价长白落叶松纸浆材优良家系。
木芯基本密度与解析木基本密度、气干密度、径面抗劈力、抗弯弹性模量、抗弯强度、顺纹抗压强度和径面硬度呈极显著正相关,且回归分析模型比较理想,可以利用胸径木芯基本密度值预测物理力学指标值,间接选择和评价长白落叶松建筑材优良家系。
4.通过生长性状、木芯基本密度和木芯管胞长宽比综合分析选出166、169为优良家系,优良家系材积、木芯基本密度和木芯管胞长宽比的遗传增益(超过对照)分别为48.34%(38.46%)、14.01%(3.68%)和19.89%(6.41%)。
5.以纸浆材为选育目标,对生长和材性、制浆造纸特性进行综合评价。选出的优良家系与木芯选择的结果一致。优良家系综纤维素含量、抗张指数和撕裂指数分别比对照高3.05%、3.74%和1.10%,早材壁腔比比对照小21.44%。
6.以建筑材为选育目标,对生长和木材物理力学性状进行综合评价。选出的优良家系,与木芯和纸浆材选择的结果一致；优良家系径面抗劈力、抗弯弹性模量和弦面硬度分别比对照高2.50%、6.90%和45.24%,早材微纤丝角、晚材率和气干体积干缩率分别比对照小2.51%、15.08%和13.30%。
In this study, the testing forest of the24-year-old Larix olgensis excellent progenies in the Jiangshanjiao experimented forest station of Heilongjiang Academy of Forestry was treated as the research object, which went through the DBH wood core and felled wood sampling methods. Then the growth traits and wood core material traits (4traits totally involved), analytic wood material traits (21traits involved), pulp and paper making properties (16traits) and the mechanics traits (20ones) of the14families went through genetic variation analysis, square variance, correlation analysis and comprehensive index analysis.
After these analyses, I obtained the genetic variation regulations of growth traits and wood material quality traits among families, researched the relations among these traits, established the regression models of wood basic density, tracheid traits and the correspondent traits of analytic wood, and finally selected out two superior families that can be utilized in both paper pulp manufacturing and construction. In addition, these results provided theories for thinning and reconstructing of vegetative seed garden of the rudimentary degree, and laid better foundations for the parent selection of hybrid breeding and the selecting of the excellent tree species of the far latter generations in the seed garden.
1. The genetic variation of traits. There are larger variations in growth traits, wood basic density, wood tracheid traits, the material traits of analytic wood, pulp and paper making traits and the mechanics intensity of the timber, and there are abundant variations among the individuals within the same family. The differences among the growth traits of each family are very significant, and the heritability values of height, breast diameter and timber volume are0.73,0.72and0.80respectively; the differences in the wood basic density and wood tracheid traits among families are significant, and the heritability values are between0.56and0.80; the differences in the micro fiber angle of earlywood of analytic wood, tracheid length, the ratio of well lo cavity of earlywood. ligin. arabinose content, tensile strength, tearing strength, etc. are very significant, and the heritability values are between0.63and0.86; the basic density of analytic wood,the differences in latewood rate, the ratio of tracheid length to width, holocellulose content, etc. are significant, and the heritability values are between0.51and0.61; the differences in cleavage strength of the tangential facet, the end-face hardness and the hardness of the tangential facet are very significant, and the heritability values are0.77,0.62and0.72respectively; the differences in drying density, latewood rate, volume shrinkage rate, the cleavage strength of the diametric facet, the MOE and the shearing strength of the diametric facet along the grain are significant, the heritability values are between0.52and0.60, and because these traits are controlled by heritability of medium and higher intensity, there is larger potential for improving them.
2. The correlation analysis among traits. The growth traits and the tracheid length are positively and very significantly correlated whose correlation relations are mainly controlled by genetic mechanisms, the two are also insignificantly correlated with wood density, the ratio of tracheid length to width and most of the mechanics intensity indices of the timber, and therefore, the two can be selected independently; wood density is significantly or very significantly and positively correlated with latewood rate, the diametric shrinkage rate, the tangential cleavage strength. MOE. bending strength, the shearing strength of the diametric facet along the grain, the crushing strength along the grain, the hardness of the end-face and the diametric facet, but insignificantly correlated with the other traits of pulpwood, which proves that the improvement of wood density is beneficial to the synthesized improvement of the mechanics traits of the timber, and both the wood density and the traits of pulpwood can be selected independently; the ratio of the trachied length to width is significantly or very significantly and positively correlated with the tangential diameter of earlywood, the tensile strength of paper, tensile index, tearing degree and tearing index, and negatively and very significantly correlated with tracheid ratio. Holocellulose is very significantly and positively correlated with the yield rate of gross pulp and fine pulp, and significantly or very significantly and negatively correlated with the content of the extracted substance from Phenylpropanol, the substance from cold water, arabinose content and the hardness of pulp. The micro fiber angle of earlywood is positively and very significantly correlated with the dual wall-thickness and negatively and significantly correlated with tracheid length, the ratio of tracheid length to width, tearing strength degree and index and MOE; latewood rate is positively and very significantly correlated with cleavage strength, MOE, bending strength, crushing strength parallel to the grain, positively and significantly correlated with the ratio of wall to cavity of earlywood, the ratio of length to width of pulp fiber and the hardness of the end-face, significantly or very significantly and negatively correlated with the diametric diameter of earlywood, the resin canal ratio, the pulp fiber width, the ratio of wall to cavity of pulpwood, tensile strength, tensile index, bursting strength and index.
3. Wood basic density, tracheid length, tracheid width and the ratio of tracheid length to width are very significantly and positively correlated with the correspondent traits of analytic wood and the regression model is rather ideal and I can take advantage of the traits of wood at the breast height to predict the values of a single plant and indirectly select and assess the superior families specialized in pulpwood.
The wood basic density is positively and significantly correlated with the basic density of analytic wood, drying density, the diametric cleave strength, MOE, bending strength, the crushing strength parallel to the grain and the hardness of the diametric facet, the regression model is rather ideal and I can take advantage of the value of the wood density at the breast height to predict the index values of mechanics and indirectly select and assess the superior families specialized in construction timber.
4. Through the synthesized analysis of growth traits, wood basic density and the ratio of wood tracheid length to width. No.166and169families were selected as the superior families, The heritability gains of the timber volume, wood basic density and the ratio of the wood tracheid length to width are48.34%,14.01%and19.89%respectively and larger than the contrast ones by38.46%,3.68%and6.41%respectively.
5. The pulpwood was treated as the breeding target, through synthesized assessment of the growth traits, material traits and pulp and paper making traits, which was in line with the results obtained according to the growth traits, material quality traits and the physical traits of paper pulp of analytic wood. The sleeted families of the holocellulose content, tensile index, tearing degree larger than the contrast ones by3.05%,3.74%and1.10%, the the ratio of well to cavity of earlywood less than the contrast ones by21.44%.
6.The construction timber was treated as the breeding target, through synthesized assessment of growth traits and mechanics traits, which was in line with the results obtained according to the material quality traits and the physical traits of paper pulp of analytic wood, the superior families of the diametric cleave strength, MOE, hardness of the tangential facet larger than the contrast ones by2.50%.6.90%and45.24%respectively, microfiber angle of early and latewood, latewood rate and volume shrinkage rate less than2.51%,15.08%and13.30%.
1.性状的遗传变异：生长性状、木芯基本密度和管胞性状、解析木材质性状和制浆造纸性状以及木材物理力学强度均存在较大的变异,家系内个体间也存在着丰富的变异。家系间生长性状差异极显著,树高、胸径和材积的家系遗传力分别为0.73、0.72和0.80。家系间木芯基本密度、木芯管胞性状差异显著,家系遗传力在0.56-0.80之间。解析木早材微纤丝角、管胞长度、早材壁腔比、木质素、阿拉伯糖含量、抗张强度、撕裂度等家系间差异极显著,家系遗传力在0.63-0.86之间,基本密度、晚材率、管胞长宽比、综纤维素含量等家系间差异显著,家系遗传力在0.51-0.61；家系间弦面抗劈力、端面硬度和弦面硬度差异极显著,家系遗传力分别为0.77、0.62和0.72；家系间气干密度、晚材率、气干体积干缩率、径面抗劈力、抗弯弹性模量和径面顺纹抗剪强度差异显著,家系遗传力在0.52-0.60之间,这些性状受中等及以上强度的遗传控制,家系水平的材性改良潜力较大
2.性状相关分析：牛长性状与管胞长度呈极显著正相关,且相关关系主要受遗传机制控制,与木材密度、管胞长宽比相关不显著,与大多数木材力学强度指标相关不显著,可以进行独立选择；木材密度与晚材率、径向干缩率、弦面抗劈力、抗弯弹性模量、抗弯强度、径面顺纹抗剪强度、顺纹抗压强度、硬度(端面、径面)呈极显著或显著正相关,与纸浆材其它性状相关不显著,说明木材密度的改良有利于木材物理力学性状的综合改良,与纸浆材性状可以独立选择；管胞长宽比与早材弦向直径、纸的抗张强度、抗张指数、撕裂度和撕裂指数呈极显著或显著正相关,与管胞比量呈极显著负相关。综纤维素含量与粗浆得率、细浆得率呈极显著正相关,与苯醇抽出物含量、冷水抽提物含量、阿拉伯糖含量和浆的硬度呈极显著或显著负相关。早材纤丝角与早材双壁厚呈极显著正相关,与管胞长度和管胞长宽比、撕裂度和撕裂指数、木材抗弯弹性模量呈显著负相关；晚材率与抗劈力、抗弯弹性模量、抗弯强度、顺纹抗压强度呈极显著正相关,与早材壁腔比、浆料纤维长宽比、木材端而硬度呈显著正相关,与早材径向直径、树脂道比量、浆料纤维宽度、浆料壁腔比、抗张强度、抗张指数、耐破度和耐破指数呈极显著或显著负相关。
3.木芯基本密度、管胞长度、管胞宽度以及管胞长宽比与解析木相应性状呈极显著正相关关系,且回归模型较为理想,可以利用胸径木芯各性状值预测其单株值,间接选择和评价长白落叶松纸浆材优良家系。
木芯基本密度与解析木基本密度、气干密度、径面抗劈力、抗弯弹性模量、抗弯强度、顺纹抗压强度和径面硬度呈极显著正相关,且回归分析模型比较理想,可以利用胸径木芯基本密度值预测物理力学指标值,间接选择和评价长白落叶松建筑材优良家系。
4.通过生长性状、木芯基本密度和木芯管胞长宽比综合分析选出166、169为优良家系,优良家系材积、木芯基本密度和木芯管胞长宽比的遗传增益(超过对照)分别为48.34%(38.46%)、14.01%(3.68%)和19.89%(6.41%)。
5.以纸浆材为选育目标,对生长和材性、制浆造纸特性进行综合评价。选出的优良家系与木芯选择的结果一致。优良家系综纤维素含量、抗张指数和撕裂指数分别比对照高3.05%、3.74%和1.10%,早材壁腔比比对照小21.44%。
6.以建筑材为选育目标,对生长和木材物理力学性状进行综合评价。选出的优良家系,与木芯和纸浆材选择的结果一致；优良家系径面抗劈力、抗弯弹性模量和弦面硬度分别比对照高2.50%、6.90%和45.24%,早材微纤丝角、晚材率和气干体积干缩率分别比对照小2.51%、15.08%和13.30%。
In this study, the testing forest of the24-year-old Larix olgensis excellent progenies in the Jiangshanjiao experimented forest station of Heilongjiang Academy of Forestry was treated as the research object, which went through the DBH wood core and felled wood sampling methods. Then the growth traits and wood core material traits (4traits totally involved), analytic wood material traits (21traits involved), pulp and paper making properties (16traits) and the mechanics traits (20ones) of the14families went through genetic variation analysis, square variance, correlation analysis and comprehensive index analysis.
After these analyses, I obtained the genetic variation regulations of growth traits and wood material quality traits among families, researched the relations among these traits, established the regression models of wood basic density, tracheid traits and the correspondent traits of analytic wood, and finally selected out two superior families that can be utilized in both paper pulp manufacturing and construction. In addition, these results provided theories for thinning and reconstructing of vegetative seed garden of the rudimentary degree, and laid better foundations for the parent selection of hybrid breeding and the selecting of the excellent tree species of the far latter generations in the seed garden.
1. The genetic variation of traits. There are larger variations in growth traits, wood basic density, wood tracheid traits, the material traits of analytic wood, pulp and paper making traits and the mechanics intensity of the timber, and there are abundant variations among the individuals within the same family. The differences among the growth traits of each family are very significant, and the heritability values of height, breast diameter and timber volume are0.73,0.72and0.80respectively; the differences in the wood basic density and wood tracheid traits among families are significant, and the heritability values are between0.56and0.80; the differences in the micro fiber angle of earlywood of analytic wood, tracheid length, the ratio of well lo cavity of earlywood. ligin. arabinose content, tensile strength, tearing strength, etc. are very significant, and the heritability values are between0.63and0.86; the basic density of analytic wood,the differences in latewood rate, the ratio of tracheid length to width, holocellulose content, etc. are significant, and the heritability values are between0.51and0.61; the differences in cleavage strength of the tangential facet, the end-face hardness and the hardness of the tangential facet are very significant, and the heritability values are0.77,0.62and0.72respectively; the differences in drying density, latewood rate, volume shrinkage rate, the cleavage strength of the diametric facet, the MOE and the shearing strength of the diametric facet along the grain are significant, the heritability values are between0.52and0.60, and because these traits are controlled by heritability of medium and higher intensity, there is larger potential for improving them.
2. The correlation analysis among traits. The growth traits and the tracheid length are positively and very significantly correlated whose correlation relations are mainly controlled by genetic mechanisms, the two are also insignificantly correlated with wood density, the ratio of tracheid length to width and most of the mechanics intensity indices of the timber, and therefore, the two can be selected independently; wood density is significantly or very significantly and positively correlated with latewood rate, the diametric shrinkage rate, the tangential cleavage strength. MOE. bending strength, the shearing strength of the diametric facet along the grain, the crushing strength along the grain, the hardness of the end-face and the diametric facet, but insignificantly correlated with the other traits of pulpwood, which proves that the improvement of wood density is beneficial to the synthesized improvement of the mechanics traits of the timber, and both the wood density and the traits of pulpwood can be selected independently; the ratio of the trachied length to width is significantly or very significantly and positively correlated with the tangential diameter of earlywood, the tensile strength of paper, tensile index, tearing degree and tearing index, and negatively and very significantly correlated with tracheid ratio. Holocellulose is very significantly and positively correlated with the yield rate of gross pulp and fine pulp, and significantly or very significantly and negatively correlated with the content of the extracted substance from Phenylpropanol, the substance from cold water, arabinose content and the hardness of pulp. The micro fiber angle of earlywood is positively and very significantly correlated with the dual wall-thickness and negatively and significantly correlated with tracheid length, the ratio of tracheid length to width, tearing strength degree and index and MOE; latewood rate is positively and very significantly correlated with cleavage strength, MOE, bending strength, crushing strength parallel to the grain, positively and significantly correlated with the ratio of wall to cavity of earlywood, the ratio of length to width of pulp fiber and the hardness of the end-face, significantly or very significantly and negatively correlated with the diametric diameter of earlywood, the resin canal ratio, the pulp fiber width, the ratio of wall to cavity of pulpwood, tensile strength, tensile index, bursting strength and index.
3. Wood basic density, tracheid length, tracheid width and the ratio of tracheid length to width are very significantly and positively correlated with the correspondent traits of analytic wood and the regression model is rather ideal and I can take advantage of the traits of wood at the breast height to predict the values of a single plant and indirectly select and assess the superior families specialized in pulpwood.
The wood basic density is positively and significantly correlated with the basic density of analytic wood, drying density, the diametric cleave strength, MOE, bending strength, the crushing strength parallel to the grain and the hardness of the diametric facet, the regression model is rather ideal and I can take advantage of the value of the wood density at the breast height to predict the index values of mechanics and indirectly select and assess the superior families specialized in construction timber.
4. Through the synthesized analysis of growth traits, wood basic density and the ratio of wood tracheid length to width. No.166and169families were selected as the superior families, The heritability gains of the timber volume, wood basic density and the ratio of the wood tracheid length to width are48.34%,14.01%and19.89%respectively and larger than the contrast ones by38.46%,3.68%and6.41%respectively.
5. The pulpwood was treated as the breeding target, through synthesized assessment of the growth traits, material traits and pulp and paper making traits, which was in line with the results obtained according to the growth traits, material quality traits and the physical traits of paper pulp of analytic wood. The sleeted families of the holocellulose content, tensile index, tearing degree larger than the contrast ones by3.05%,3.74%and1.10%, the the ratio of well to cavity of earlywood less than the contrast ones by21.44%.
6.The construction timber was treated as the breeding target, through synthesized assessment of growth traits and mechanics traits, which was in line with the results obtained according to the material quality traits and the physical traits of paper pulp of analytic wood, the superior families of the diametric cleave strength, MOE, hardness of the tangential facet larger than the contrast ones by2.50%.6.90%and45.24%respectively, microfiber angle of early and latewood, latewood rate and volume shrinkage rate less than2.51%,15.08%and13.30%.
引文
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