爆裂玉米主基因—多基因遗传体系及遗传多样性研究
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摘要
本试验通过爆裂玉米遗传差异较大的二个纯系亲本组合B01(P_1)×B02(P_2)6个家系世代(P_1、P_2、F_1、B_(1：2)、B_(2：2)、F_(2：3))，采用重复内分组随机设计，研究爆裂玉米主要数量性状的主基因-多基因遗传体系及选用45份不同来源的爆裂玉米自交系用SSR分子标记方法对遗传多样性进行分析，为掌握爆裂玉米数量性状遗传规律，提高爆裂玉米遗传育种效率，提供理论依据。所获得主要结果如下：
1．爆花率性状遗传体系由2对主基因+多基因所组成。两对主基因存在加性、显性效应，但差异较大。同时两对主基因间存在加×加、加×显和显×显等互作效应。爆花率除受主基因控制外，也受多基因控制。多基因效应主要为加性和显性、上位性效应。B_(1：2)、B_(2：2)、F_(2：3)家系分离世代主基因遗传率(h_(mg)~2(％))分别为74.99％、78.35％、62.33％；多基因遗传率(h_(pg)~2(％))分别为3.12％、3.52％、6.12％。可见三个家系世代的表型变异的60％多由主基因决定。环境效应对爆花率也有一定的影响。
2．膨胀倍数性状遗传体系由一对主基因和多基因所组成。主基因存在加性效应，无显性效应。多基因存在加性、显性效应两者均为负值。B_(1：2)、B_(2：2)、F_(2：3)家系分离世代主基因遗传率(h_(mg)~2(％))均较小，B_(2：2)家系世代多基因遗传率为89.16％，所以，多基因对膨胀倍数性状表型变异起主要作用。
3．膨胀体积性状遗传体系由多基因所组成。多基因存在加性、显性、上位性效应，多基因遗传率在10.49％—65.52％。环境效应对膨胀体积表型变异有较大的影响。
4．淀粉含量和蛋白质含量性状遗传体系由一对主基因+多基因所组成。淀粉含量主基因存在加性效应，无显性效应存在。主基因的加性效应d=-10.0643，为负值，表明主基因存在时，淀粉含量降低，多基因存在加性、显性效应，但显性效应为负值。B_(2：2)、F_(2：3)世代主基因遗传率分别为80.96％、82.55％较高，多基因遗传率为2.91％、3.27％。淀粉含量表型变异主要由主基因决定的；蛋白质含量主基因存在完全显性效应。显性效应为负值；多基因存在加性、显性效应，显性效应大于加性效应，B_(1：2)、F_(2：3)世代主基因遗传率分别为50.16％、52.80％，即表型变异的50％多由主基因决定的。环境对蛋白质含量表型变异有一定的影响。
5．百粒重、单穗重、行粒数、株高性状遗传体系均由一对主基因和多基因所组成(D-2)。主基因存在加性效应，无显性效应存在。百粒重、株高性状主基因加性效应为正值，多基因存在加性、显性效应均为正值，因此2个性状存在杂种优势。百粒重F_(2：3)，家系世代主基因遗传率为70.33％，多基因遗传率为12.81％；株高F_(2：3)家系世代主基因遗传率为53.87％，多基因遗传率为38.54％，百粒重、株高的表型变异主要由主基因决定的。单穗重、行粒数性状主基因加性效应为负值，单穗重多基因加性、显性效应均为正值；行粒数多基因加性效应为负值，显性效应为正值，显性效应大于加性效应。单穗重B_(2：2)世代主基因遗传率为62.11％，多基因遗传率为32.48％，单穗重表型变异主要由主基因决定的，环境变异影响较小；行粒数B_(2：2)世代主基因遗传率为48.31％，多基因遗传率为22.19％，环境效应对行粒数影响较大。
6．穗长、穗位高、雄穗分枝数性状遗传体系由多基因所组成。多基因存在加性、显性、上位性。多基因遗传率较大，穗长、穗位高、雄穗分枝数F_(2：3)群体多基因遗传率分别为88.52％、78.12％、84.82％。
7．秃尖长、雄穗长度、叶绿素含量性状遗传体系由一对主基因+多基因所组成。秃尖长主基因存在加性、显性，加性效应为负值，显性效应为正值，而且加性效应大于显性效应，多基因存在加性、显性，加性效应为正值，显性效应为负值。显性度h／d为-0.1058，表明有秃尖对无秃尖为显性。B_(2：2)世代主基因遗传率为69.05％，多基因遗传率为27.29％，环境效应对秃尖长影响较小；雄穗长度主基因存在完全显性，显性效应为负值。多基因存在加性、显性。显性效应大于加性效应。B_(2：2)主基因遗传率为58.88％，多基因遗传率为33.96％。雄穗长度的表型变异主基因起重要作用，环境影响较小；叶绿素含量主基因存在加性、显性，加性效应为负值，显性效应为正值，而且显性效应大于加性效应，显性度为-2.0688。多基因存在加性、显性、上位性。B_(2：2)世代主基因遗传率为55.28％，多基因遗传率为26.36％。叶绿素含量表型变异主要由主基因决定的。环境效应也有一定的影响。
8．选用的80对SSR引物在45份爆裂玉米自交系间共检测出334个等位基因变异，每对引物检测出2-9个等位基因，平均为4.2个，多态性信息量变化范围为0.10-0.83，平均为0.53。
9．将45份爆裂玉米自交系划分为5类群，第Ⅰ类群包括B01、B36、B45、B17、B10、B28、B39、B38等8个自交系；第Ⅱ类群包括B02、B41、B14、B43、B07、B13、B40、B03、B20、B22、B33、B35、B16、B25、B04、B15、B23、B37、B21、B42、B11、B24、B05、B30、B32、B19、B44、B08、B27、B12、B26、B29、B31等33个自交系；第Ⅲ类群包括B09自交系；第Ⅳ类群包括B34自交系；第Ⅴ类群包括B06、B18等2个自交系。
10．在一定范围内，亲本自交系间遗传距离越大，杂种优势越强，因此在育种实践中，应该把自交系分子遗传距离作为重要参考。从本试验看几乎所有强优势组合均为不同类群自交系间杂交种，但也并非类群间自交系所组配的组合一定是强优势组合。因此，利用分子标记的遗传距离不足以有效预测供试爆裂玉米自交系间的杂种优势。
11．爆裂玉米小区产量、一般配合力效应与分子标记遗传距离呈显著正相关关系，F_1爆花率、膨胀倍数与分子标记遗传距离之间无显著的相关关系。
Major gene-polygene genetic system of popcorn quantitative trait were studied byduplicational grouping random design in 2 pureline as parent combination B01 (P_1)×B02(P_2),6ancestry generation, and genetic diversity were studied by SSR molecular marker in 45popcorns.The purpose of these works is to hold the genetic law of popcorn quantitative trait,enhance breeding efficiency.The results is below:
1. The trait genetic system of popcorn flower is compose of 2 pair major gene whichexhibit additive and dominance effect and large different, and polygene, meanwhile, 2pair exist additive×additive, additive×dominance and dominance×dominanceinteractive effect. Beside manin gene, the popcorn flower rate is controled by polygene.Polygne effect main express additive, dominance and interactive effect. The major geneheritability(h_(mg)~2(%)) of ancestry segregate generation of B_(1:2), B_(2:2), F_(2:3) is74.99%,78.35%and 62.33%, polyene heritability(h_(pg)~2(%)) is 3.12%,3.52%,6.12%respectly. The 60% phenotype variation of 3 ancestory generation is controled by majorgene, and a small quantity by environment.
2. Expand trait genetic system is compose 1 pair major gene and polygene. Major genewxhibit additive effect, no dominance effect. Polygene exist additive and dominanceeffect, and all is negative value. The major gene heritability of B_(1:2), B_(2:2), F_(2:3) are lower,the polygene heritability of B_(2:2) is 89.16%, so, expand trait is main controled bypolygene.
3. Expand volumn trait is compose of polygene. Polygene exhibit additive, dominance,interactive effect, polygene heritability range from 10.49%-65.52%. Environmentinfluence expand volumn phenotype.
4. Starch conent and protein content trait is compose by lpair major gene and polygene.Starch content major gene exist additive effect, no dominance effect. Additive effect ofmajor gene d-10.0643, is negative, that indicate major gene exist, the starch content islower, polygene exist additive, dominance effect, which is negative value. The majorgene heritability of B_(2:2), F_(2:3) is 80.96%, 82.55%, the heritability of polygene is 2.91%, 3.27%. The phenotype variance is due to gene, the major gene of proein exhibitcomplete dominance effect which is negative value. The polygene exist additive,dominance effect, and later is larger than before. The major gene heritability of B_(1:2), F_(2:3)generation is 50.16%, 52.80%, i.e.major gene determine phenotype variance about 50%,and environment has some influence.
5. 100-grains weight, single spike weight, grain number per row and stem high geneticsystem is compose 1 pair major gene and polygene. Major gene exist additive effect, nodominance effect. 100-grains weight,stem high trait major gene additive effect ispositive, polygene exist additive, dominance effect all positive, hence the 2 characterexist heterosis. 100-grain weight heritability of F_(2:3) major gene is 70.33%, theheritability of polygene is 12.81%, the stem high major gene heritability of F_(2:3) ancestrygeneration is 53.87%, polygene heritability is 38.54%, the major gene play importantrole in 100-grains weight, stem high. The additive effect of major gene grain number perrow is negative, polygene additive, dominance effect of a single spike weight is positive,polygene additive effect of grain number per row is negative, dominance effect ispositive, dominance effect is more than additive effect. A single spike weight major geneheritability of B_(2:2) is 62.11%, polygene is 32.48%, a single spike phenotype variancemain due to mainene, a little environment, the major gene heritability of B_(2:2) grainnumber per row is 48.31%, polygeneis 22.19%, the environment play important role inthe trait.
6. Spike length, the position of spike, male spike branch genetic systemis composepolygene. Polygene exist additive,dominance, interactive effect. Polygen heritability islarger, spike length, the heritability of position of spike and male spike branch in F_(2:3) isrespectively 88.52%,78.12%, 84.82%.
7. Bare tip long, male spike long, chlorophyll content genetic system is compose 1 pairand polygene. Major gene of bare tip long has additive, dominance effect, the former isnegative, the later is positive, and former is larger, than later, polygene exist additive,dominance effect, and the former is positive, the later is negative. The dominance degreeis -0.1056, that showed that bare tip is dominance than no. the major gene heritability ofB_(2:2) is 69.05%, polygene is 27.29%, the environment affect bare tip little. The majorgene of male spike long exist complete dominance, effect is negative. Polygene includeadditive, dominance, the later large former. The major gene heritability of B_(2:2) is 58.88%, polygene heritability is 33.96%. the gene play important role in the trait of male spikelong, and a little environment, the major gene of chlorophyll content include additive,dominance, the former is negative, the later is positive, and dominance is larger additive,the dominance degree is -2.0688. polygene include additive, dominance and interactive.The major gene heritability of B_(2:2) is 55.28%, polygene heritability is 26.36%. the geneplay important role in the trait of chlorophyll content, and some environment.
8. 334 alleles variance were checked by using 80 pair SSR primer, each primer check 2-9allele, average 4.2, polymorohic variance from 0.10 to 0.83, average 0.53.
9. 45 popcorn inbred lines group 5, the first group includeB01,B36,B45,B17,B10,B28,B39,B38; the second include B02, B41, B14, B43, B07,B13, B40, B03, B20, B22, B33, B35, B16, B25, B04, B15, B23, B37, B21, B42, B11,B24, B05, B30, B32, B19, B44, B08, B27, B12, B26, B29, B31; the third include B09;the forth include B34; the fifth include B06, B18.
10. In some content, the genetic distance more large, the heterosis is strength, and it isimportant reference in breeding. Almost all strength heterosis is express by differentclasses inbred line, but not all combinance from different classes express like so. Hence,molecule marker is not enough to estimate heterosis of popcorn inbred line.
11. The plot yield, general combining ability and molecule marker exhibit positivecorrelation, popcornflower rate of F_1, expand and molecule marker exhibit no dominantcorrelation.
1．爆花率性状遗传体系由2对主基因+多基因所组成。两对主基因存在加性、显性效应，但差异较大。同时两对主基因间存在加×加、加×显和显×显等互作效应。爆花率除受主基因控制外，也受多基因控制。多基因效应主要为加性和显性、上位性效应。B_(1：2)、B_(2：2)、F_(2：3)家系分离世代主基因遗传率(h_(mg)~2(％))分别为74.99％、78.35％、62.33％；多基因遗传率(h_(pg)~2(％))分别为3.12％、3.52％、6.12％。可见三个家系世代的表型变异的60％多由主基因决定。环境效应对爆花率也有一定的影响。
2．膨胀倍数性状遗传体系由一对主基因和多基因所组成。主基因存在加性效应，无显性效应。多基因存在加性、显性效应两者均为负值。B_(1：2)、B_(2：2)、F_(2：3)家系分离世代主基因遗传率(h_(mg)~2(％))均较小，B_(2：2)家系世代多基因遗传率为89.16％，所以，多基因对膨胀倍数性状表型变异起主要作用。
3．膨胀体积性状遗传体系由多基因所组成。多基因存在加性、显性、上位性效应，多基因遗传率在10.49％—65.52％。环境效应对膨胀体积表型变异有较大的影响。
4．淀粉含量和蛋白质含量性状遗传体系由一对主基因+多基因所组成。淀粉含量主基因存在加性效应，无显性效应存在。主基因的加性效应d=-10.0643，为负值，表明主基因存在时，淀粉含量降低，多基因存在加性、显性效应，但显性效应为负值。B_(2：2)、F_(2：3)世代主基因遗传率分别为80.96％、82.55％较高，多基因遗传率为2.91％、3.27％。淀粉含量表型变异主要由主基因决定的；蛋白质含量主基因存在完全显性效应。显性效应为负值；多基因存在加性、显性效应，显性效应大于加性效应，B_(1：2)、F_(2：3)世代主基因遗传率分别为50.16％、52.80％，即表型变异的50％多由主基因决定的。环境对蛋白质含量表型变异有一定的影响。
5．百粒重、单穗重、行粒数、株高性状遗传体系均由一对主基因和多基因所组成(D-2)。主基因存在加性效应，无显性效应存在。百粒重、株高性状主基因加性效应为正值，多基因存在加性、显性效应均为正值，因此2个性状存在杂种优势。百粒重F_(2：3)，家系世代主基因遗传率为70.33％，多基因遗传率为12.81％；株高F_(2：3)家系世代主基因遗传率为53.87％，多基因遗传率为38.54％，百粒重、株高的表型变异主要由主基因决定的。单穗重、行粒数性状主基因加性效应为负值，单穗重多基因加性、显性效应均为正值；行粒数多基因加性效应为负值，显性效应为正值，显性效应大于加性效应。单穗重B_(2：2)世代主基因遗传率为62.11％，多基因遗传率为32.48％，单穗重表型变异主要由主基因决定的，环境变异影响较小；行粒数B_(2：2)世代主基因遗传率为48.31％，多基因遗传率为22.19％，环境效应对行粒数影响较大。
6．穗长、穗位高、雄穗分枝数性状遗传体系由多基因所组成。多基因存在加性、显性、上位性。多基因遗传率较大，穗长、穗位高、雄穗分枝数F_(2：3)群体多基因遗传率分别为88.52％、78.12％、84.82％。
7．秃尖长、雄穗长度、叶绿素含量性状遗传体系由一对主基因+多基因所组成。秃尖长主基因存在加性、显性，加性效应为负值，显性效应为正值，而且加性效应大于显性效应，多基因存在加性、显性，加性效应为正值，显性效应为负值。显性度h／d为-0.1058，表明有秃尖对无秃尖为显性。B_(2：2)世代主基因遗传率为69.05％，多基因遗传率为27.29％，环境效应对秃尖长影响较小；雄穗长度主基因存在完全显性，显性效应为负值。多基因存在加性、显性。显性效应大于加性效应。B_(2：2)主基因遗传率为58.88％，多基因遗传率为33.96％。雄穗长度的表型变异主基因起重要作用，环境影响较小；叶绿素含量主基因存在加性、显性，加性效应为负值，显性效应为正值，而且显性效应大于加性效应，显性度为-2.0688。多基因存在加性、显性、上位性。B_(2：2)世代主基因遗传率为55.28％，多基因遗传率为26.36％。叶绿素含量表型变异主要由主基因决定的。环境效应也有一定的影响。
8．选用的80对SSR引物在45份爆裂玉米自交系间共检测出334个等位基因变异，每对引物检测出2-9个等位基因，平均为4.2个，多态性信息量变化范围为0.10-0.83，平均为0.53。
9．将45份爆裂玉米自交系划分为5类群，第Ⅰ类群包括B01、B36、B45、B17、B10、B28、B39、B38等8个自交系；第Ⅱ类群包括B02、B41、B14、B43、B07、B13、B40、B03、B20、B22、B33、B35、B16、B25、B04、B15、B23、B37、B21、B42、B11、B24、B05、B30、B32、B19、B44、B08、B27、B12、B26、B29、B31等33个自交系；第Ⅲ类群包括B09自交系；第Ⅳ类群包括B34自交系；第Ⅴ类群包括B06、B18等2个自交系。
10．在一定范围内，亲本自交系间遗传距离越大，杂种优势越强，因此在育种实践中，应该把自交系分子遗传距离作为重要参考。从本试验看几乎所有强优势组合均为不同类群自交系间杂交种，但也并非类群间自交系所组配的组合一定是强优势组合。因此，利用分子标记的遗传距离不足以有效预测供试爆裂玉米自交系间的杂种优势。
11．爆裂玉米小区产量、一般配合力效应与分子标记遗传距离呈显著正相关关系，F_1爆花率、膨胀倍数与分子标记遗传距离之间无显著的相关关系。
Major gene-polygene genetic system of popcorn quantitative trait were studied byduplicational grouping random design in 2 pureline as parent combination B01 (P_1)×B02(P_2),6ancestry generation, and genetic diversity were studied by SSR molecular marker in 45popcorns.The purpose of these works is to hold the genetic law of popcorn quantitative trait,enhance breeding efficiency.The results is below:
1. The trait genetic system of popcorn flower is compose of 2 pair major gene whichexhibit additive and dominance effect and large different, and polygene, meanwhile, 2pair exist additive×additive, additive×dominance and dominance×dominanceinteractive effect. Beside manin gene, the popcorn flower rate is controled by polygene.Polygne effect main express additive, dominance and interactive effect. The major geneheritability(h_(mg)~2(%)) of ancestry segregate generation of B_(1:2), B_(2:2), F_(2:3) is74.99%,78.35%and 62.33%, polyene heritability(h_(pg)~2(%)) is 3.12%,3.52%,6.12%respectly. The 60% phenotype variation of 3 ancestory generation is controled by majorgene, and a small quantity by environment.
2. Expand trait genetic system is compose 1 pair major gene and polygene. Major genewxhibit additive effect, no dominance effect. Polygene exist additive and dominanceeffect, and all is negative value. The major gene heritability of B_(1:2), B_(2:2), F_(2:3) are lower,the polygene heritability of B_(2:2) is 89.16%, so, expand trait is main controled bypolygene.
3. Expand volumn trait is compose of polygene. Polygene exhibit additive, dominance,interactive effect, polygene heritability range from 10.49%-65.52%. Environmentinfluence expand volumn phenotype.
4. Starch conent and protein content trait is compose by lpair major gene and polygene.Starch content major gene exist additive effect, no dominance effect. Additive effect ofmajor gene d-10.0643, is negative, that indicate major gene exist, the starch content islower, polygene exist additive, dominance effect, which is negative value. The majorgene heritability of B_(2:2), F_(2:3) is 80.96%, 82.55%, the heritability of polygene is 2.91%, 3.27%. The phenotype variance is due to gene, the major gene of proein exhibitcomplete dominance effect which is negative value. The polygene exist additive,dominance effect, and later is larger than before. The major gene heritability of B_(1:2), F_(2:3)generation is 50.16%, 52.80%, i.e.major gene determine phenotype variance about 50%,and environment has some influence.
5. 100-grains weight, single spike weight, grain number per row and stem high geneticsystem is compose 1 pair major gene and polygene. Major gene exist additive effect, nodominance effect. 100-grains weight,stem high trait major gene additive effect ispositive, polygene exist additive, dominance effect all positive, hence the 2 characterexist heterosis. 100-grain weight heritability of F_(2:3) major gene is 70.33%, theheritability of polygene is 12.81%, the stem high major gene heritability of F_(2:3) ancestrygeneration is 53.87%, polygene heritability is 38.54%, the major gene play importantrole in 100-grains weight, stem high. The additive effect of major gene grain number perrow is negative, polygene additive, dominance effect of a single spike weight is positive,polygene additive effect of grain number per row is negative, dominance effect ispositive, dominance effect is more than additive effect. A single spike weight major geneheritability of B_(2:2) is 62.11%, polygene is 32.48%, a single spike phenotype variancemain due to mainene, a little environment, the major gene heritability of B_(2:2) grainnumber per row is 48.31%, polygeneis 22.19%, the environment play important role inthe trait.
6. Spike length, the position of spike, male spike branch genetic systemis composepolygene. Polygene exist additive,dominance, interactive effect. Polygen heritability islarger, spike length, the heritability of position of spike and male spike branch in F_(2:3) isrespectively 88.52%,78.12%, 84.82%.
7. Bare tip long, male spike long, chlorophyll content genetic system is compose 1 pairand polygene. Major gene of bare tip long has additive, dominance effect, the former isnegative, the later is positive, and former is larger, than later, polygene exist additive,dominance effect, and the former is positive, the later is negative. The dominance degreeis -0.1056, that showed that bare tip is dominance than no. the major gene heritability ofB_(2:2) is 69.05%, polygene is 27.29%, the environment affect bare tip little. The majorgene of male spike long exist complete dominance, effect is negative. Polygene includeadditive, dominance, the later large former. The major gene heritability of B_(2:2) is 58.88%, polygene heritability is 33.96%. the gene play important role in the trait of male spikelong, and a little environment, the major gene of chlorophyll content include additive,dominance, the former is negative, the later is positive, and dominance is larger additive,the dominance degree is -2.0688. polygene include additive, dominance and interactive.The major gene heritability of B_(2:2) is 55.28%, polygene heritability is 26.36%. the geneplay important role in the trait of chlorophyll content, and some environment.
8. 334 alleles variance were checked by using 80 pair SSR primer, each primer check 2-9allele, average 4.2, polymorohic variance from 0.10 to 0.83, average 0.53.
9. 45 popcorn inbred lines group 5, the first group includeB01,B36,B45,B17,B10,B28,B39,B38; the second include B02, B41, B14, B43, B07,B13, B40, B03, B20, B22, B33, B35, B16, B25, B04, B15, B23, B37, B21, B42, B11,B24, B05, B30, B32, B19, B44, B08, B27, B12, B26, B29, B31; the third include B09;the forth include B34; the fifth include B06, B18.
10. In some content, the genetic distance more large, the heterosis is strength, and it isimportant reference in breeding. Almost all strength heterosis is express by differentclasses inbred line, but not all combinance from different classes express like so. Hence,molecule marker is not enough to estimate heterosis of popcorn inbred line.
11. The plot yield, general combining ability and molecule marker exhibit positivecorrelation, popcornflower rate of F_1, expand and molecule marker exhibit no dominantcorrelation.
引文
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