滇楸优树及其子代苗期性状遗传变异研究
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摘要
为选育适应于西南高山地区栽培的良种，探究滇楸（Catalpa fargesii Bur. f. duclouxii(Dode) Gilmour）的群体遗传多样性、优良种源、家系和无性系的选育，在贵州、云南滇楸资源集中分布的区域，进行滇楸群体遗传多样性的取样调查，对优良的种质进行选择收集评价、对优树进行苗期子代测定，按照良种选育程序通过遗传参数估算选育优良的家系和无性系。通过以上研究获得以下主要研究结果：
一、滇楸群体表型性状的变异类型非常丰富，遗传多样性水平高。17个种群花部的表型性状在种群间及种群内差异均显著。种群间变异系数为19.88%～35.75%，性状平均表型分化系数（VST）为28.26%，分化幅度为18.91%～42.15%，表型频率分化系数（Pst）为0.5147，Shannon-Weaver多样性指数值（H'）在1.3490～2.5225之间，滇楸花部表型性状的重复力高，最高值花下萼唇长平均为0.685，花性状中花上萼唇长、下萼唇长和花径性状的稳定性强。花部性状与地理变异有明显的相关性，其中花上萼唇长、下萼唇长、花径与纬度呈极显著相关，花枝长度与经度显著相关并与海拔呈极显著负相关。
滇楸果实8个表型性状在滇楸的种群间及种群内均表现极显著差异。种群间的变异系数为31.03%（24.53%～39.28%），性状表型分化系数VST分别为22.81%（10.53%～39.28%），果实平均多样度系数Lst、Ls和Pst分别为0.0359、0.755和0.0453，Shannon-Weaver多样性指数（H'）在1.6184～2.3617之间。果实性状间重复力平均值为0.543，变化幅度在0.399～0.694之间。相对花的重复力较低。对来自不同省份的9个种群果实性状进行UPGMA聚类，结果表明滇楸果实的表型变异并没有依地理距离而聚类，与花部性状聚类结果一致。
二、滇楸不同气候区、种源、家系种子发芽性状遗传变异。滇楸的种子质量和播种品质在气候区、种源、家系间及家系内性状间变异系数的差异十分明显。其种子千粒重、场圃发芽率、发芽势、发芽指数等4个性状变异系数的变化范围分别为2.41%～20.57%、3.67%～28.89%、8.92%～30.82%和8.75%～34.13%之间。种子千粒重与7月均温和有效积温均显著的负相关，相关系数R值为-0.593和-0.662。
三、滇楸家系子代苗期生长性状变异十分丰富。一、二年生苗木的苗高、地径等性状方差分析显示，在气候区类型、种源、家系三个层次上的差异显著。一年生苗高、地径差异均达到极显著水平，二年生苗性状在不同家系间除苗高、地径在气候类型区间差异不显著外，其余的差异均达到极显著水平。苗期生长性状的相关分析表明，一、二年生地径生长与产地七月均温均呈显著相关（R=0.672,和R=0.595），二年生地径与经度呈显著负相关（R=-0.589），苗高与地径生长量在一、二年均呈极显著正相关。一年生苗高、地径的家系遗传力分别为0.805和0.664，二年生苗高、地径的家系遗传力分别为0.837和0.603。一、二年生家系间苗高、地径的遗传变异系数分别为18.798%、11.053%和10.141%、6.589%；一、二年生家系间苗高、地径的表型变异系数分别为39.477%、28.461%和28.818%、35.408%；一二年生的遗传增益最高分别为24.96%和35.32%。对二年生苗木按I值选出最优家系31个，他们分别来源于兴仁、盘县和瓮安等10个种源。
四、滇楸优良无性系的选择。无性系生长性状、叶部性状、木材材性性状在无性系间差异显著。无性系高、叶柄长、叶面积、基本密度、胞壁率、纤维实际长和双壁厚性状的重复力分别为0.921、0.960、0.946、0.842、0.872、0.792和0.725。无性系木材基本密度与解剖性状胞壁率、双壁厚、弦向胞腔直径、弦向中央直径分别显著相关。纤维实际长与各性状间相关不显著。通过遗传参数、相关分析、主成分、聚类分析和综合评分方法等，生长性状与叶部性状结合，筛选出7个优良无性系；联合生长、木材解剖与力学性状，选出生长量大，解剖结构好，木材物理性状稳定的5个优良无性系。
For selecting and breeding improved varieties that are suitable for cultivate in southwestmountain, genetic diversity of populations, heredity and variation of germplasm provenances,and families and clones breeding are studied in Catalpa fargesii Bur. f. duclouxii (Dode)Gilmour. In according to the procedure to estimate the superior families by genetic parameters,in Guizhou and Yunnan where Catalpa fargesii distribute concentrately, genetic diversity ofpopulation were investigated, excellent germplasm were selected, collected and evaluated, aswell as progeny test of the collecting excellent tree were conducted. The above researchobtains main results as following:
Firstly, variation of phenotypic traits was vary rich, and its genetic diversity was high levelin population of Catalpa fargesii. Floral phenotypic differences were significantly big within17populations and among17populations. Variation coefficient among population is19.88%～35.75%, the mean phenotypic differentiation coefficient of characters (VST) was28.26%, therange of change was18.91%～42.15%. Differentiating coefficient of phenotype frequency(Pst) is0.5147. Shannon-Weaver diversity index (H') varies was1.3490～2.5225. Repeatabilityof floral phenotypic traits in catalpa fargesii was high, its highest value was the flower calyx,namely0.685on average. Morethan, length of flower calyx lip, length of calyx lip anddiameter of flower traits has strong stability. There was a significant correlation between floraltraits and geographic factors, and length of flower calyx lip, length of calyx lip, flowerdiameter was significantly correlated with the latitude and longitude, squid length wassignificant negatively correlated with altitude.8fruit phenotypic traits differentiatesignificantly among populations and within populations in Catalpa fargesii. The variationcoefficient among populations was31.03%(24.53%～39.28%), Phenotypic differentiationcoefficient VSTwas22.81%(10.53%～39.28%), Lst, Ls and Pst of the average fruit diversitycoefficient were0.0359,0.755and0.0453respectively, Shannon-Weaver diversity index (H') was in1.6184～2.3617. Mean value of repeatability in the fruit characters was0.543, variationrange was in0.399～0.694, it was lower than flower characters. By UPGMA cluster of fruitcharacters in9populations from different provinces, the result showed that the fruit phenotypicvariation in Catalpa fargesii was not in accordance with the geographical distance, but was inaccordance with the results of floral traits cluster.
Secondly, seed germinating variation of different climate, provenance, family in Catalpafargesii. Variation coefficient of seed quality and sowing quality in Catalpa fargesii has veryobvious difference in climate, provenances, families. Range of variation coefficient in the seedweight, germination rate, germination potential, and germination index were2.41%～20.57%,3.67%～28.89%,8.92%～30.82%and8.75%～34.13%respectively. Seed weight weresignificantly negative correlation with mean tempreture in July and effective accumulatedtemperature, R value of the correlation coefficient was-0.593and-0.662.
Thirdly, growth traits variating of family progeny was very rich in Catalpa fargesii. Byvariance analysis for one or biennial s, traits of the height and ground diameter showsignificant differences in climate types, provenance and family. Difference of height andground diameter for one year old reached significant level, and2year old height andground diameter of different families had no significant difference in climate types, but otherdifference reached significant level. Correlation analysis of showed that one or biennialdiameter growth were significant (R=0.672and0.595) with July temperature in origin. thebiennial diameter showed a significant negative correlation with longitude (R=-0.589).Family heritability of height, ground diameter for one year old were0.805and0.664respectively. Family heritability of height, ground diameter for two year old were0.837and0.603respectively. One or biennial family genetic variation coefficients of height and grounddiameter were18.798%,11.053%and10.141%,6.589%respectively; the phenotypic variationcoefficients of height and ground diameter of family for one or biennial year were39.477%,28.461%and28.818%,35.408%. Genetic gain of one or two years was up to24.96%and35.32%respectively. Acording to I value,31superior families were selected from seedings for two year old, that distributed in10provenances, eg: XingRen county, Pan county and Wengancounty so on.
Fourthly, superior clones selecting in Catalpa fargesii. There were significant differencesin growth traits, leaf characters and wood properties of clone among clones. Repeatability ofheight, footstalk length, leave area, basic density, cell wall percentage, fiber length and doublewall thickness were respectively0.921,0.960,0.946,0.842,0.872,0.792and0.725in clones.Wood basic density of clones were significantly related to anatomical characters of plant cellwall, double wall thickness, ratio of diameter to chord, cellular tangential diameter. It was notsignificant correlation between actual fiber length and other traits. According to the geneticparameters, correlation analysis results, physical mechanics indexes, superior clones ingrowing rapidly, good anatomical structure and stable physical properties of wood werecollected during early selection of clones. By genetic parameter, correlating analysis, study ofprincipal component, cluster analysis, and method of synthetical score so on,7superior cloneswere selected using growth traits and leaf characters. Combining growing traits withanatomical structure and mechanics traits of wood,5superior clones were selected that weregrowing rapidly, anatomical structure well, steady physical behavior of wood.
一、滇楸群体表型性状的变异类型非常丰富，遗传多样性水平高。17个种群花部的表型性状在种群间及种群内差异均显著。种群间变异系数为19.88%～35.75%，性状平均表型分化系数（VST）为28.26%，分化幅度为18.91%～42.15%，表型频率分化系数（Pst）为0.5147，Shannon-Weaver多样性指数值（H'）在1.3490～2.5225之间，滇楸花部表型性状的重复力高，最高值花下萼唇长平均为0.685，花性状中花上萼唇长、下萼唇长和花径性状的稳定性强。花部性状与地理变异有明显的相关性，其中花上萼唇长、下萼唇长、花径与纬度呈极显著相关，花枝长度与经度显著相关并与海拔呈极显著负相关。
滇楸果实8个表型性状在滇楸的种群间及种群内均表现极显著差异。种群间的变异系数为31.03%（24.53%～39.28%），性状表型分化系数VST分别为22.81%（10.53%～39.28%），果实平均多样度系数Lst、Ls和Pst分别为0.0359、0.755和0.0453，Shannon-Weaver多样性指数（H'）在1.6184～2.3617之间。果实性状间重复力平均值为0.543，变化幅度在0.399～0.694之间。相对花的重复力较低。对来自不同省份的9个种群果实性状进行UPGMA聚类，结果表明滇楸果实的表型变异并没有依地理距离而聚类，与花部性状聚类结果一致。
二、滇楸不同气候区、种源、家系种子发芽性状遗传变异。滇楸的种子质量和播种品质在气候区、种源、家系间及家系内性状间变异系数的差异十分明显。其种子千粒重、场圃发芽率、发芽势、发芽指数等4个性状变异系数的变化范围分别为2.41%～20.57%、3.67%～28.89%、8.92%～30.82%和8.75%～34.13%之间。种子千粒重与7月均温和有效积温均显著的负相关，相关系数R值为-0.593和-0.662。
三、滇楸家系子代苗期生长性状变异十分丰富。一、二年生苗木的苗高、地径等性状方差分析显示，在气候区类型、种源、家系三个层次上的差异显著。一年生苗高、地径差异均达到极显著水平，二年生苗性状在不同家系间除苗高、地径在气候类型区间差异不显著外，其余的差异均达到极显著水平。苗期生长性状的相关分析表明，一、二年生地径生长与产地七月均温均呈显著相关（R=0.672,和R=0.595），二年生地径与经度呈显著负相关（R=-0.589），苗高与地径生长量在一、二年均呈极显著正相关。一年生苗高、地径的家系遗传力分别为0.805和0.664，二年生苗高、地径的家系遗传力分别为0.837和0.603。一、二年生家系间苗高、地径的遗传变异系数分别为18.798%、11.053%和10.141%、6.589%；一、二年生家系间苗高、地径的表型变异系数分别为39.477%、28.461%和28.818%、35.408%；一二年生的遗传增益最高分别为24.96%和35.32%。对二年生苗木按I值选出最优家系31个，他们分别来源于兴仁、盘县和瓮安等10个种源。
四、滇楸优良无性系的选择。无性系生长性状、叶部性状、木材材性性状在无性系间差异显著。无性系高、叶柄长、叶面积、基本密度、胞壁率、纤维实际长和双壁厚性状的重复力分别为0.921、0.960、0.946、0.842、0.872、0.792和0.725。无性系木材基本密度与解剖性状胞壁率、双壁厚、弦向胞腔直径、弦向中央直径分别显著相关。纤维实际长与各性状间相关不显著。通过遗传参数、相关分析、主成分、聚类分析和综合评分方法等，生长性状与叶部性状结合，筛选出7个优良无性系；联合生长、木材解剖与力学性状，选出生长量大，解剖结构好，木材物理性状稳定的5个优良无性系。
For selecting and breeding improved varieties that are suitable for cultivate in southwestmountain, genetic diversity of populations, heredity and variation of germplasm provenances,and families and clones breeding are studied in Catalpa fargesii Bur. f. duclouxii (Dode)Gilmour. In according to the procedure to estimate the superior families by genetic parameters,in Guizhou and Yunnan where Catalpa fargesii distribute concentrately, genetic diversity ofpopulation were investigated, excellent germplasm were selected, collected and evaluated, aswell as progeny test of the collecting excellent tree were conducted. The above researchobtains main results as following:
Firstly, variation of phenotypic traits was vary rich, and its genetic diversity was high levelin population of Catalpa fargesii. Floral phenotypic differences were significantly big within17populations and among17populations. Variation coefficient among population is19.88%～35.75%, the mean phenotypic differentiation coefficient of characters (VST) was28.26%, therange of change was18.91%～42.15%. Differentiating coefficient of phenotype frequency(Pst) is0.5147. Shannon-Weaver diversity index (H') varies was1.3490～2.5225. Repeatabilityof floral phenotypic traits in catalpa fargesii was high, its highest value was the flower calyx,namely0.685on average. Morethan, length of flower calyx lip, length of calyx lip anddiameter of flower traits has strong stability. There was a significant correlation between floraltraits and geographic factors, and length of flower calyx lip, length of calyx lip, flowerdiameter was significantly correlated with the latitude and longitude, squid length wassignificant negatively correlated with altitude.8fruit phenotypic traits differentiatesignificantly among populations and within populations in Catalpa fargesii. The variationcoefficient among populations was31.03%(24.53%～39.28%), Phenotypic differentiationcoefficient VSTwas22.81%(10.53%～39.28%), Lst, Ls and Pst of the average fruit diversitycoefficient were0.0359,0.755and0.0453respectively, Shannon-Weaver diversity index (H') was in1.6184～2.3617. Mean value of repeatability in the fruit characters was0.543, variationrange was in0.399～0.694, it was lower than flower characters. By UPGMA cluster of fruitcharacters in9populations from different provinces, the result showed that the fruit phenotypicvariation in Catalpa fargesii was not in accordance with the geographical distance, but was inaccordance with the results of floral traits cluster.
Secondly, seed germinating variation of different climate, provenance, family in Catalpafargesii. Variation coefficient of seed quality and sowing quality in Catalpa fargesii has veryobvious difference in climate, provenances, families. Range of variation coefficient in the seedweight, germination rate, germination potential, and germination index were2.41%～20.57%,3.67%～28.89%,8.92%～30.82%and8.75%～34.13%respectively. Seed weight weresignificantly negative correlation with mean tempreture in July and effective accumulatedtemperature, R value of the correlation coefficient was-0.593and-0.662.
Thirdly, growth traits variating of family progeny was very rich in Catalpa fargesii. Byvariance analysis for one or biennial s, traits of the height and ground diameter showsignificant differences in climate types, provenance and family. Difference of height andground diameter for one year old reached significant level, and2year old height andground diameter of different families had no significant difference in climate types, but otherdifference reached significant level. Correlation analysis of showed that one or biennialdiameter growth were significant (R=0.672and0.595) with July temperature in origin. thebiennial diameter showed a significant negative correlation with longitude (R=-0.589).Family heritability of height, ground diameter for one year old were0.805and0.664respectively. Family heritability of height, ground diameter for two year old were0.837and0.603respectively. One or biennial family genetic variation coefficients of height and grounddiameter were18.798%,11.053%and10.141%,6.589%respectively; the phenotypic variationcoefficients of height and ground diameter of family for one or biennial year were39.477%,28.461%and28.818%,35.408%. Genetic gain of one or two years was up to24.96%and35.32%respectively. Acording to I value,31superior families were selected from seedings for two year old, that distributed in10provenances, eg: XingRen county, Pan county and Wengancounty so on.
Fourthly, superior clones selecting in Catalpa fargesii. There were significant differencesin growth traits, leaf characters and wood properties of clone among clones. Repeatability ofheight, footstalk length, leave area, basic density, cell wall percentage, fiber length and doublewall thickness were respectively0.921,0.960,0.946,0.842,0.872,0.792and0.725in clones.Wood basic density of clones were significantly related to anatomical characters of plant cellwall, double wall thickness, ratio of diameter to chord, cellular tangential diameter. It was notsignificant correlation between actual fiber length and other traits. According to the geneticparameters, correlation analysis results, physical mechanics indexes, superior clones ingrowing rapidly, good anatomical structure and stable physical properties of wood werecollected during early selection of clones. By genetic parameter, correlating analysis, study ofprincipal component, cluster analysis, and method of synthetical score so on,7superior cloneswere selected using growth traits and leaf characters. Combining growing traits withanatomical structure and mechanics traits of wood,5superior clones were selected that weregrowing rapidly, anatomical structure well, steady physical behavior of wood.
引文
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