水稻籼粳亚种间杂交衍生系遗传基础的研究
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摘要
水稻籼粳亚种间杂种F1蕴藏着巨大的杂种优势。有效利用水稻籼粳亚种间杂种优势是全球产稻国稻作科技的一个热点。从育种实践的角度,创制水稻籼粳亚种间杂交衍生系谱、揭示其遗传基础,对培育实用型强优势籼粳亚种间杂交稻具有重要的理论意义与指导价值。本文在评述水稻籼粳亚种间杂交性状遗传育种、籼粳亚种分类、遗传多样性和亲缘关系研究进展的基础上,以一个历时21年4个育种阶段,涉及18个亲本和39个不同阶段育成的籼粳交衍生系谱为研究材料,从分子水平上揭示其籼粳分化与杂种优势关系、遗传多样性与系谱亲缘关系;从遗传生态学角度分析不同育种阶段衍生系产量相关性状的改良效果、育成品系产量相关性状的遗传及其与环境互作效应、评价其育种利用价值,旨在为进一步有效利用水稻籼粳杂交衍生系及其杂交稻配组提供科学依据。主要结果如下:
1、采用58个籼粳特异ILP分子标记结合程氏形态指数法,检测了包括18个亲本、39个衍生系和2个对照品种的水稻籼粳亚种间杂交衍生系谱的籼粳分化度。以籼粳属性判别值(Di)和籼(粳)稻成分指数值分别作为判别不同籼粳类型多态性条带和籼粳类型划分的定性指标。发现供试材料基因组DNA在所检测位点上均存在籼粳分化,不存在100%纯籼(或粳)类型;18个亲本中共检测出4个粳型、5个偏粳型、8个偏籼型和1个籼型,平均粳稻成分指数分别为79.69%、32.08%、10.94%和6.25%;39个籼粳杂交衍生系中共检测出1个粳型、11个偏粳型、20个偏籼型和7个籼型,平均粳稻成分指数分别为68.70%、24.62%、12.92%和4.76%;对照品种日本晴和9311分别属于粳型和籼型,粳稻成分指数分别为95.83 %和4.17 %。同时,发现供试材料外观形态上属于粳型的C418和明恢502,其粳稻成分分别仅有31.25%(偏粳型)和10.42%(偏籼型);而属于籼型的9308和明恢63,其粳稻成分分别为33.33%(偏粳型)和12.50%(偏籼型)。进一步分析认为以明恢63为亲本配组的杂交稻汕优63之所以雄居全球推广面积之首,其原因之一是利用了部分籼粳亚种间杂种优势。另外,籼(或粳)稻成分指数分别与程氏形态指数多数指标间表现明显的相关关系,且籼粳类型的划分与形态指数判别的结果均表现相同的趋势,表明采用ILP分子标记获得的籼(或粳)稲成分指数及定性指标作为判别籼粳分化类型是合理的,对有效利用籼粳亚种间优良基因具有重要作用。
2、采用分布于水稻基因组的221个SSR位点结合系谱分析法,研究了水稻籼粳亚种间杂交衍生系和亲本的遗传多样性和亲缘关系。发现反映遗传多样性的多态性位点数、多态性位点百分率(PPB)、观察等位基因值(Na)、有效等位基因值(Ne)、平均Nei遗传多样性指数(H)、Shannon多样性指数(I)等指标,亲本均大于衍生系,衍生四代大于衍生二代,四种籼粳分化类型大小依次为:偏粳型﹥偏籼型﹥粳型﹥籼型;通过SSR标记可将衍生系和亲本聚为5大类,其中第三大类又可分为四个亚类,发现各类间的籼粳分化类型和亲缘关系差异明显,具有明恢63亲缘关系的均聚在第三大类第二亚类,它与其它类群存在一定的遗传距离,由此表明,多数的衍生系在拓宽三系恢复系遗传基础上有明显改进,采用Structure群体遗传结构法也得出类似的结果;主成分分析(PCA)结果发现随着衍生系的不断改良,籼稻遗传成分逐渐增加,衍生系中籼(粳)成分比例更趋于合理。系谱追踪等位变异法(PAM)分析亲本遗传物质在衍生世代间的传递,发现ketanNangka、嘉南粳、02428、明恢72等4个祖先亲本对衍生一代92gk729的遗传贡献率分别为15.09%,21.32%,22.78%和34.64%,对衍生二代的分别为6.05%,7.09%,7.98%和13.41%,对衍生三代的分别为3.51%,4.04%,4.56%和7.92%,对衍生四代的分别为3.31%,3.23%,3.68%和7.55%,祖先亲本对各衍生世代的遗传贡献率大小依次为明恢72>02428>嘉南粳>Ketan Nangka ;各级亲本及杂交重组后的特异遗传位点也间接地传递给衍生后代,对衍生系的遗传贡献是随着衍生代数的递增而逐渐下降。值得一提的是,92gk729有10个经祖先亲本遗传重组后保留的SSR特异位点,其中的5个位点能稳定地分别传递到23.21%、35.71%、82.14%、64.29%、41.07%的各衍生世代品系中,平均有2.21个位点可遗传到衍生四代。采用系谱追踪等位变异法对4个祖先亲本及其衍生系分别进行聚类,发现Ketan Nangka、嘉南粳和02428三个祖先亲本及其衍生系的聚类结果较为相似,与明恢72为祖先亲本的有明显差异,较好地体现出各衍生系不同的遗传背景,有助于了解衍生系的遗传结构分化。
3、选用4个籼(粳)稻不育系和54个水稻籼粳亚种间杂交衍生系及其亲本,按不完全双列杂交(NC-Ⅱ设计)配制的216个杂交组合,探讨双亲籼粳遗传差异与产量相关性状杂种优势的关系;选用8个三系和两系不育系及其籼粳分化具有明显差异的12个衍生系,按不完全双列杂交(NC-Ⅱ设计)配制的96个杂交组合,研究双亲籼粳遗传差异的适合范围;用41个ILP标记对供试不育系进行籼粳分化检测。发现三系不育系春江12A、广抗13A、K17A、中9A、金23A和II-32A粳稻成分指数值分别为92.50%(粳)、9.76%(偏籼)、9.76%(偏籼)、5%(籼),7.32%(籼)、7.69%(籼);两系不育系N5088S、培矮64 S、广占63S、SE21S粳稻成分指数值分别为95.00%(粳)、24.39%(偏粳)、12.50%(偏籼)、7.32%(籼)。双亲籼粳遗传差异对产量相关性状杂种优势的影响因研究材料的不同有异,不同地点表现较为相似的趋势;双亲籼粳遗传差异对每穗实粒数和结实率超亲优势具有较大的影响,而对有效穗、千粒重的影响较小。双亲籼粳遗传差异适合范围的研究结果表明,三系杂交稻双亲粳稻成分指数差值(恢复系-不育系)在-5%时,杂种的单株谷重性状表现最小的负向优势;处于-5%～0之间,随着差值的减少,正向优势逐渐增强;在0～20%之间,随着差值的扩大,正向优势均大于20%;在20%～25%之间,正向优势趋于下降。当差值分别在0～10%和10%～20%范围内,恢复系恰好为偏籼和偏粳类型,均可筛选出杂种优势较强的杂交稻。两系杂交稻组合的双亲籼粳遗传差异的适合范围比三系杂交组合更宽,在7.7%～15%差值范围内较易选出强优组合。
4、在双亲籼粳遗传差异适合范围研究材料的基础上,进一步分析了双亲籼粳遗传差异对茎秆和光合性状杂种优势的影响。结果表明,在所考查的杂交稻组合中,双亲粳稻成分指数差值与倒伏指数中亲优势呈显著负相关;与抗折力、茎壁截面积的超亲和中亲优势均呈显著或极显著正相关;与弯曲力矩和茎椭圆截面积的杂种优势相关不明显;其中,三系杂交组合的抗折力、茎壁截面积的超亲优势与双亲粳稻成分指数差值均呈明显的正相关关系。在所考查的杂交稻组合中,双亲粳稻成分指数差值与齐穗期光合速率的超亲(中亲)优势均呈显著的负相关关系,而在成熟期则表现正相关趋势。
5、选用不同衍生世代的8个水稻籼粳亚种间杂交品系和8个三系和两系不育系,按不完全双列杂交(NC-Ⅱ设计)配制包括亲本和杂种一代两个世代的遗传材料,采用基因型与环境互作的加性-显性遗传模型及相应的统计方法,分析了不同衍生世代9个产量相关性状的表型值、遗传效应和杂种优势的变化趋势。表型值分析表明,亲本的单株谷重随着衍生世代的增加呈明显增重的趋势,F1在衍生二代和衍生四代变化都不明显,但两者均比衍生一代有较大提高;亲本和F1的每穗总粒数和每穗实粒数都随着衍生世代的增加呈明显增多的趋势,株高在衍生二代和衍生四代变化都不明显,但两者均比衍生一代降高效果显著,抽穗期在衍生世代之间变化幅度不大,表明在控制F1生育期超亲上有明显的改良效果。遗传效应分析认为,每穗总粒数和每穗实粒数的加性方差占表型方差的比率在衍生世代间均呈增强趋势,而显性方差占表型方差的比率则呈下降趋势,环境影响的效应变化不明显;生育期和千粒重的加性方差占表型方差的比率在衍生四代均大于衍生一代和衍生二代,而显性方差占表型方差的比率则最低,环境影响的效应变化不明显。衍生四代的上述性状受显性效应影响较小,通过选择能稳定遗传。杂种优势分析可见,在遗传主效应和环境效应的群体平均优势和群体超亲优势上,生育期在衍生世代间的变化不明显,多数为负向效应,表现缩短生育期的趋势;单株有效穗数在衍生世代间有增加的趋势,但多数还是为负向优势;每穗总粒数、每穗实粒数和株高在衍生世代间有降低趋势,但多数为正效应。
6、选用衍生四代的8个育成品系和8个三系和两系不育系,按不完全双列杂交(NC-Ⅱ设计)配制包括亲本和杂种一代两个世代的遗传材料,采用基因型与环境互作的加性-显性遗传模型及相应的统计方法,分析了不同环境条件下育成品系产量相关性状的遗传效应和杂种优势,评价了育成品系的育种利用价值。结果表明,除单株谷重主要受基因型×环境互作效应控制(占总遗传方差的69%)外,其余8个性状主要受遗传主效应的控制,遗传主效应方差占总遗传方差的幅度为54.9%~91.3%。在遗传主效应中,抽穗期、单株有效穗、穗总粒数、穗实粒数和千粒重等性状都是以基因的加性效应为主,其加性方差分别占遗传主效应方差的76.8%、70.9%、94.3%、84.9%和85.8%,这5个性状可以通过选择能稳定遗传;单株谷重、结实率、株高和穗长等性状以基因的显性效应为主,其显性方差分别占遗传主效应方差的68.8%、78.8%、56.3%和61.3%,该4个性状可以通过杂种优势加以利用。杂种优势分析可见,除单株有效穗和抽穗期外,其余7个性状均表现显著或极显著的正向群体平均优势,单株谷重最大,达18.5%,千粒重最小,仅为3.2%。单株有效穗、千粒重和株高3个性状群体平均优势的稳定性较好;其余6个性状的稳定性较差。除单株产谷重、株高和穗长外,其余6个性状均表现明显的负向群体超亲优势。株高、抽穗期和单株有效穗的群体超亲优势稳定性较好。其余6个性状的群体超亲优势稳定性较差。育成品系育种利用价值分析表明,明恢413具有增加杂种后代每穗总粒数、穗长和单株谷重的效应;明恢509可增加杂种后代的每穗总粒数、结实率、穗长和单株谷重;明恢417对增加后代的结实率、千粒重,穗长、单株谷重具有明显作用;明恢118有提高结实率、增加单株谷重的效应;明恢502具有增长穗长,增多每穗总粒数和提高结实率的作用;明恢503和明恢512均可增加每穗总粒数;明恢398能降低株高和提高千粒重。
Huge heterosis exists in rice Indica-japonica subspecies hybrids F1. How to utilize the heterosis in indica-japonica subspecies hybrids rice efficiently is a topic of general interest in the field of hybrid rice breeding in the world. In view of the breeding practice, deriving pedigree of indica-japonica subspecies hybrids rice and its genetic basis are of theoretical importance and guiding significance in breeding for elite indica-japonica subspecies hybrids rice. In this paper, 18 parents and 39 deriving pedigrees were used to reveal the relationship between indica-japonica differentiation and heterosis, and the relationship between heredity diversity and pedigree. The improving effects of yield traits in different breeding stages and genetic effects of yield traits and their interaction by environment were also analyzed. The result would provide scientific foundation for further effective utilization of deriving pedigree of indica-japonica subspecies hybrids rice and hybrid rice combination. The main experimental results are as follows.
1. Differentiation of indica-japonica rice subspecies hybrids deriving pedigree, including 18 parents, 39 deriving pedigrees and 2 control varieties, were determinated by using 58 ILP markers (Intron Length Polymorphism Markers in rice) and Cheng’s Index method. The discrimination value Di and component index of Indica-japonica were selected as qualitative index to distinguish the amplified fragment polymorphism and classify types of indica-japonica. The results show that indica-japonica differentiation existed at the detection sites in the genomic DNA from materials. no indica type (or indica type) was detected with 100% component index. Of 18 parents, 4 japonica, 5 japonica-clinious, 8 indica-clinous and 1 indica were detected to contain 79.69%, 32.08%, 10.94%, and 6.25% of average component indexes of japonica respectively. In 39 deriving pedigrees of rice subspecies indica-japonica, 1 japonica, 11 japonica-clinious, 20 indica-clinous and 7 indica were detected to have 68.70%, 24.62%, 12.92%, and 4.76% of average component indexes of japonica respectively. The component indexes of Nippornbare and 9311, which belong to japonica and indica, were 95.83% and 4.17% respectively. The indexes of japonica rice cultivar C418 and Minghui 502 were 31.25% and 10.42% respectively, but indica cultivar 9308 and Minghui 63 were 33.33% and 12.50% respectively. Further analysis indicated that cultivar Shanyou 63 hybrid, which were derived from interbred from cultivar Minghui 63, had the largest plantation area in the world the heterosis between rice subspecies indica and japonica. There were also significant correlations between component index and Cheng’s index, and the classification of indica and japonica was similar to the dermination of shape index. Therefore, it’s reasonable to use component and determination index of indica or japonica obtained from ILP markers, in order to determinate the differentiation of indica and japonica. It’s vital to utilize the useful genes from rice subspecies indica-japonica.
2. Studies on genetic diversity and genetic relatives of indica-japonica subspecies rice hybrid deriving pedigrees and the parents was conducted by using 221 SSR and pedigree analysis. The results showed that the indexes of genetic diversity, such as polymorphic marker number, polymorphic marker percentage (PPB), effective number of alleles (Ne), observed number of alleles (Na), Shannon diversity index (H) et al., from parents were higher than those from deriving pedigrees, the forth generation deriving pedigree was higher than the second pedigree. And the order of indica and japonica differentiation from high to low is japonica-clinious, indica-clinious, japonica, indica. The deriving pedigrees and parents were divided into 5 clusters by SSR markers, and then the third cluster divided into 4 sub-clusters. There was a significant difference between different indica-japonica types of deriving pedigrees or parents. The cultivars, which were closely relative to cultivar Minghui 63, were gathered at the second sub-cluster, and the third cluster. It’s suggested that most deriving pedigrees could significantly improve the genetic basis of three line restorer. The results of population genetic structure indicates that with the improvement of deriving pedigrees, genetic component was increased and the component ratio of indica or japonica deriving pedigrees tends to be reasonable. Pedigree allele method was used to analyze the transfer of genetic materials from parents in deriving generation. The results showed that the genetic contribution rations of ancestors, such as cultivar Ketan Nangka, Jiananjin, 02428 and Minghui 72, to the first deriving generation were 15.09%, 21.32%, 22.78% and 34.36% respectively, the second generation were 6.05%, 7.09%, 7.98% and 13.41%, respectively, for the third were 3.51%, 4.04%, 4.56% and 7.92% respectively, for the forth were 3.31%, 3.23%, 3.68% and 7.55% respectively. The order of genetic contribution rations of ancestor parents to deriving generations from large to small is cultivar Minghui 72, 02428, cultivar Jiananjin, Ketan Nangka. Each parent including the specific genetic loci was also passed to the deriving offspings, and the genetic contribution to deriving pedigrees decreased with the increase of the deriving generations. Cultivar 92gk729 kept 10 SSR specific loci from ancestor parents after genetic reconstruction, the 5 loci of which could be stably passed to deriving generations with 23.21%, 35.71%, 82.14%, 64.29% and 41.07%. The forth deriving generation inherited 2.21 loci from parents on average. The cluster results of cultivar Ketan Nangka, Jiananjin and 02428 were similar. Parents and their deriving pedigrees, but had a significant difference from cultivar Minghui 72. The results suggested that the different genetic background of deriving pedigrees and helped to understand the genetic structure differentiation of deriving pedigrees.
3. Using 4 indica or japonica sterile line and 54 deriving pedigrees and their parents, 216 cross combinations were generated with incomplete diallel cross design (NC-II), so that the relation between genetic differences of biparental indica-japonica and heterosis of yield characters could be discussed. 96 cross combinations, from 8 three-line and tow-line CMS and 12 rice subspecies differentiation with significant differences, were selected in order to research on the application sphere of genetic differences of parental incica-japonica. 41 ILP markers were used to detect the indica-japonica differentiation. The results revealed that the proportion index of japonica of CMS lines cultivar Chuenjiang 12A, Guangkang 13A, K17A, Zhong 9A, Jin 23A and II-32A was 92.50% (japonica), 9.76% (indica-clinious), 9.76% (indica-clinious), 5% (indica), 7.32% (indica), 7.69% (indica), respectively. The proportion index of japonica of TGMS lines cultivar Peiai 64S, Guangzhan 63S, and SE21S were 24.39 % ( japonica-clinious), 12.50 % ( indica-clinious) and 7.32 % (indica), respectively. Effects of genetic differences of parental indica-japonica on the heterosis of yield traits were diverse for the different materials, but had same trends in different places. Genetic differences of parental indica-japonica had considerable influence on the transgressive heterosis of setting grains per panicle and setting grain percentage, but little influence on the effective panicles and 1000-grain weight. The result of research on the application sphere of genetic differences of parental incica-japonica showed that the character of grain weight per plant was the lowest negative heterosis when the difference of restorer to sterile lines was -5%. When the difference from -5% to 0, the positive heterosis increased, When the difference from 0 to 20%, the positive heterosis was much more than 20%, but the positive heterosis decreased from 20% to 25%. The hybrid rice with strong heterosis could be screened from the restorer lines, which were indica-clinious and japonica-clinious type, when the difference from 0 to 10% and 10% to 20%, respectively. The application range of genetic difference of the parental indica-japonica in two-line hybrid rice was much wider than in three-line hybrid rice. It could be easy to select high yield combinations from the difference of 7.7% to 15%.
4. On the basis of application range, the effects of genetic difference from parental indica-japonica on the heterosis of stalk and photosynthetic characters were further analyzed. The results indicated that there was a significantly or extremely negative correlation between the component index differences of parents and lodging index of heterosis, but no significant correlation between bending moment and stem ellipse section area of heterosis in all cross combinations. There was a significant correlation between the component index differences of parents with breaking resistance and stem wall section area of transgressive heterosis. The component index of parents was significantly correlated with the photosynthetic characters at full heading stage of tansgreesive or mid-parent heterosis, but at maturity there was a positive correlation.
5. By using 8 hybrid lines of subspecies rice and 8 three-line and tow-line sterile lines, two generations of parent and F1 hybrid were selected by using incomplete diallel cross design (NC-Ⅱ), so that the variation trends of phenotypic value of 9 yield characters, genetic effects and heterosis in different deriving pedigrees was analyzed with additive-dominance genetic model. The results of phenotypic value showed that grain weight per plant increased along with the increase of deriving generations. And the change of the second and the forth deriving generation was not significant, but both of them increased much more than the first deriving generation. The total seed per panicles and the setting seeds per panicles of parents and F1 aggrandized, but plant height in the second deriving pedigrees and the third deriving pedigrees changed little, but increased much more than the first deriving generation. The small change of heading stage between deriving generations indicates that transgressive heterosis has good improving effect on controlling growth period of F1. Genetic effect analysis reckoned that additive variance of total seeds per panicles and setting seeds per panicles increased of phenotypic variance, but dominant variance decreased, and the environmental effect was not obvious. The additive variance of growth period and 1000-seeds weight of phenotypic variance in the forth deriving generation was larger than those in the first and second deriving generation, and dominant variance of phenotypic variance was low. The results of heterosis analysis showed that growth period of population average and transgressive heterosis changed less in different deriving generations, and most were negative heterosis. The total seeds per panicles, setting seeds per panicles and plant height decreased, and most were positive heterosis.
6. By using 8 hybrid strains of the forth deriving generation and 8 three-line and tow-line sterile lines, two generations of parent and F1 hybrid were selected with incomplete diallel cross design (NC-Ⅱ), so that the genetic effects and heterosis of yield characters under different types of environments was analyzed by using additive-dominance genetic model, and the value in use of hybrid strains was estimated. The results show that 8 characters were controlled by the main genetic effect, and the main genetic effect variance was from 54.9% to 91.3% of genetic variance except for grain weight per plant. The main genetic effect variance of characters, such as heading stage, effective panicles per plant, total seeds per panicles, setting seeds per panicles and 1000-seeds weight, mainly was the additive variance, whose main genetic effects were 76.8%, 70.9%, 94.3%, 84.95 and 85.8%, respectively. The characters of them could be stably inherited by selection. The main genetic effect variance of characters, such as grain weight per plant, setting percentage, plant height and panicle length, mainly was the dominant variance, which was 68.8%, 78.8%, 56.3% and 61.3% of main genetic effect, respectively. These 4 characters could be utilized by heterosis. The results of heterosis analysis showed that there were significant or extremely significant positive heterosis in 7 characters, except effective panicles per plant and heading stage. The grain weight per plant was the biggest (18.5%), and 1000-grain weight was the smallest, only 3.2%. The stability of population average heterosis of effective panicles, 1000-grain weight and plant height was better than the other 6 characters. There were significant negative population transgressive heterosis in the 6 characters, except for grain weight per plant, plant height and panicle length. The stability of population trangressive heterosis of plant height, heading stage and effective panicle per plant was better than other 6 characters. Analysis of breeding value for breed strains indicated that cultivar Minghui 413 could increase effect for the total seeds per panicles, panicles length and grain weight per plant in hybrid progeny, cultivar Minghui 509 could increase effect for the total seeds per panicles, seeds setting rate, panicles length and grain weight per plant, cultivar Minghui 417 could increase effect for setting percentage, 1000-grain weight, panicles length and grain weight per plant, cultivar Minghui 118 could increase effect for setting percentage and grain weight per plant, cultivar Minghui 502 could increase effect for panicles length, total seeds per panicles and seeds setting rate, cultivar Minghui 503 and 512 could increase effect for total seeds per panicles, Minghui 398 could decrease effect for plant height and increase effect for 1000- grain weight.
1、采用58个籼粳特异ILP分子标记结合程氏形态指数法,检测了包括18个亲本、39个衍生系和2个对照品种的水稻籼粳亚种间杂交衍生系谱的籼粳分化度。以籼粳属性判别值(Di)和籼(粳)稻成分指数值分别作为判别不同籼粳类型多态性条带和籼粳类型划分的定性指标。发现供试材料基因组DNA在所检测位点上均存在籼粳分化,不存在100%纯籼(或粳)类型;18个亲本中共检测出4个粳型、5个偏粳型、8个偏籼型和1个籼型,平均粳稻成分指数分别为79.69%、32.08%、10.94%和6.25%;39个籼粳杂交衍生系中共检测出1个粳型、11个偏粳型、20个偏籼型和7个籼型,平均粳稻成分指数分别为68.70%、24.62%、12.92%和4.76%;对照品种日本晴和9311分别属于粳型和籼型,粳稻成分指数分别为95.83 %和4.17 %。同时,发现供试材料外观形态上属于粳型的C418和明恢502,其粳稻成分分别仅有31.25%(偏粳型)和10.42%(偏籼型);而属于籼型的9308和明恢63,其粳稻成分分别为33.33%(偏粳型)和12.50%(偏籼型)。进一步分析认为以明恢63为亲本配组的杂交稻汕优63之所以雄居全球推广面积之首,其原因之一是利用了部分籼粳亚种间杂种优势。另外,籼(或粳)稻成分指数分别与程氏形态指数多数指标间表现明显的相关关系,且籼粳类型的划分与形态指数判别的结果均表现相同的趋势,表明采用ILP分子标记获得的籼(或粳)稲成分指数及定性指标作为判别籼粳分化类型是合理的,对有效利用籼粳亚种间优良基因具有重要作用。
2、采用分布于水稻基因组的221个SSR位点结合系谱分析法,研究了水稻籼粳亚种间杂交衍生系和亲本的遗传多样性和亲缘关系。发现反映遗传多样性的多态性位点数、多态性位点百分率(PPB)、观察等位基因值(Na)、有效等位基因值(Ne)、平均Nei遗传多样性指数(H)、Shannon多样性指数(I)等指标,亲本均大于衍生系,衍生四代大于衍生二代,四种籼粳分化类型大小依次为:偏粳型﹥偏籼型﹥粳型﹥籼型;通过SSR标记可将衍生系和亲本聚为5大类,其中第三大类又可分为四个亚类,发现各类间的籼粳分化类型和亲缘关系差异明显,具有明恢63亲缘关系的均聚在第三大类第二亚类,它与其它类群存在一定的遗传距离,由此表明,多数的衍生系在拓宽三系恢复系遗传基础上有明显改进,采用Structure群体遗传结构法也得出类似的结果;主成分分析(PCA)结果发现随着衍生系的不断改良,籼稻遗传成分逐渐增加,衍生系中籼(粳)成分比例更趋于合理。系谱追踪等位变异法(PAM)分析亲本遗传物质在衍生世代间的传递,发现ketanNangka、嘉南粳、02428、明恢72等4个祖先亲本对衍生一代92gk729的遗传贡献率分别为15.09%,21.32%,22.78%和34.64%,对衍生二代的分别为6.05%,7.09%,7.98%和13.41%,对衍生三代的分别为3.51%,4.04%,4.56%和7.92%,对衍生四代的分别为3.31%,3.23%,3.68%和7.55%,祖先亲本对各衍生世代的遗传贡献率大小依次为明恢72>02428>嘉南粳>Ketan Nangka ;各级亲本及杂交重组后的特异遗传位点也间接地传递给衍生后代,对衍生系的遗传贡献是随着衍生代数的递增而逐渐下降。值得一提的是,92gk729有10个经祖先亲本遗传重组后保留的SSR特异位点,其中的5个位点能稳定地分别传递到23.21%、35.71%、82.14%、64.29%、41.07%的各衍生世代品系中,平均有2.21个位点可遗传到衍生四代。采用系谱追踪等位变异法对4个祖先亲本及其衍生系分别进行聚类,发现Ketan Nangka、嘉南粳和02428三个祖先亲本及其衍生系的聚类结果较为相似,与明恢72为祖先亲本的有明显差异,较好地体现出各衍生系不同的遗传背景,有助于了解衍生系的遗传结构分化。
3、选用4个籼(粳)稻不育系和54个水稻籼粳亚种间杂交衍生系及其亲本,按不完全双列杂交(NC-Ⅱ设计)配制的216个杂交组合,探讨双亲籼粳遗传差异与产量相关性状杂种优势的关系;选用8个三系和两系不育系及其籼粳分化具有明显差异的12个衍生系,按不完全双列杂交(NC-Ⅱ设计)配制的96个杂交组合,研究双亲籼粳遗传差异的适合范围;用41个ILP标记对供试不育系进行籼粳分化检测。发现三系不育系春江12A、广抗13A、K17A、中9A、金23A和II-32A粳稻成分指数值分别为92.50%(粳)、9.76%(偏籼)、9.76%(偏籼)、5%(籼),7.32%(籼)、7.69%(籼);两系不育系N5088S、培矮64 S、广占63S、SE21S粳稻成分指数值分别为95.00%(粳)、24.39%(偏粳)、12.50%(偏籼)、7.32%(籼)。双亲籼粳遗传差异对产量相关性状杂种优势的影响因研究材料的不同有异,不同地点表现较为相似的趋势;双亲籼粳遗传差异对每穗实粒数和结实率超亲优势具有较大的影响,而对有效穗、千粒重的影响较小。双亲籼粳遗传差异适合范围的研究结果表明,三系杂交稻双亲粳稻成分指数差值(恢复系-不育系)在-5%时,杂种的单株谷重性状表现最小的负向优势;处于-5%～0之间,随着差值的减少,正向优势逐渐增强;在0～20%之间,随着差值的扩大,正向优势均大于20%;在20%～25%之间,正向优势趋于下降。当差值分别在0～10%和10%～20%范围内,恢复系恰好为偏籼和偏粳类型,均可筛选出杂种优势较强的杂交稻。两系杂交稻组合的双亲籼粳遗传差异的适合范围比三系杂交组合更宽,在7.7%～15%差值范围内较易选出强优组合。
4、在双亲籼粳遗传差异适合范围研究材料的基础上,进一步分析了双亲籼粳遗传差异对茎秆和光合性状杂种优势的影响。结果表明,在所考查的杂交稻组合中,双亲粳稻成分指数差值与倒伏指数中亲优势呈显著负相关;与抗折力、茎壁截面积的超亲和中亲优势均呈显著或极显著正相关;与弯曲力矩和茎椭圆截面积的杂种优势相关不明显;其中,三系杂交组合的抗折力、茎壁截面积的超亲优势与双亲粳稻成分指数差值均呈明显的正相关关系。在所考查的杂交稻组合中,双亲粳稻成分指数差值与齐穗期光合速率的超亲(中亲)优势均呈显著的负相关关系,而在成熟期则表现正相关趋势。
5、选用不同衍生世代的8个水稻籼粳亚种间杂交品系和8个三系和两系不育系,按不完全双列杂交(NC-Ⅱ设计)配制包括亲本和杂种一代两个世代的遗传材料,采用基因型与环境互作的加性-显性遗传模型及相应的统计方法,分析了不同衍生世代9个产量相关性状的表型值、遗传效应和杂种优势的变化趋势。表型值分析表明,亲本的单株谷重随着衍生世代的增加呈明显增重的趋势,F1在衍生二代和衍生四代变化都不明显,但两者均比衍生一代有较大提高;亲本和F1的每穗总粒数和每穗实粒数都随着衍生世代的增加呈明显增多的趋势,株高在衍生二代和衍生四代变化都不明显,但两者均比衍生一代降高效果显著,抽穗期在衍生世代之间变化幅度不大,表明在控制F1生育期超亲上有明显的改良效果。遗传效应分析认为,每穗总粒数和每穗实粒数的加性方差占表型方差的比率在衍生世代间均呈增强趋势,而显性方差占表型方差的比率则呈下降趋势,环境影响的效应变化不明显;生育期和千粒重的加性方差占表型方差的比率在衍生四代均大于衍生一代和衍生二代,而显性方差占表型方差的比率则最低,环境影响的效应变化不明显。衍生四代的上述性状受显性效应影响较小,通过选择能稳定遗传。杂种优势分析可见,在遗传主效应和环境效应的群体平均优势和群体超亲优势上,生育期在衍生世代间的变化不明显,多数为负向效应,表现缩短生育期的趋势;单株有效穗数在衍生世代间有增加的趋势,但多数还是为负向优势;每穗总粒数、每穗实粒数和株高在衍生世代间有降低趋势,但多数为正效应。
6、选用衍生四代的8个育成品系和8个三系和两系不育系,按不完全双列杂交(NC-Ⅱ设计)配制包括亲本和杂种一代两个世代的遗传材料,采用基因型与环境互作的加性-显性遗传模型及相应的统计方法,分析了不同环境条件下育成品系产量相关性状的遗传效应和杂种优势,评价了育成品系的育种利用价值。结果表明,除单株谷重主要受基因型×环境互作效应控制(占总遗传方差的69%)外,其余8个性状主要受遗传主效应的控制,遗传主效应方差占总遗传方差的幅度为54.9%~91.3%。在遗传主效应中,抽穗期、单株有效穗、穗总粒数、穗实粒数和千粒重等性状都是以基因的加性效应为主,其加性方差分别占遗传主效应方差的76.8%、70.9%、94.3%、84.9%和85.8%,这5个性状可以通过选择能稳定遗传;单株谷重、结实率、株高和穗长等性状以基因的显性效应为主,其显性方差分别占遗传主效应方差的68.8%、78.8%、56.3%和61.3%,该4个性状可以通过杂种优势加以利用。杂种优势分析可见,除单株有效穗和抽穗期外,其余7个性状均表现显著或极显著的正向群体平均优势,单株谷重最大,达18.5%,千粒重最小,仅为3.2%。单株有效穗、千粒重和株高3个性状群体平均优势的稳定性较好;其余6个性状的稳定性较差。除单株产谷重、株高和穗长外,其余6个性状均表现明显的负向群体超亲优势。株高、抽穗期和单株有效穗的群体超亲优势稳定性较好。其余6个性状的群体超亲优势稳定性较差。育成品系育种利用价值分析表明,明恢413具有增加杂种后代每穗总粒数、穗长和单株谷重的效应;明恢509可增加杂种后代的每穗总粒数、结实率、穗长和单株谷重;明恢417对增加后代的结实率、千粒重,穗长、单株谷重具有明显作用;明恢118有提高结实率、增加单株谷重的效应;明恢502具有增长穗长,增多每穗总粒数和提高结实率的作用;明恢503和明恢512均可增加每穗总粒数;明恢398能降低株高和提高千粒重。
Huge heterosis exists in rice Indica-japonica subspecies hybrids F1. How to utilize the heterosis in indica-japonica subspecies hybrids rice efficiently is a topic of general interest in the field of hybrid rice breeding in the world. In view of the breeding practice, deriving pedigree of indica-japonica subspecies hybrids rice and its genetic basis are of theoretical importance and guiding significance in breeding for elite indica-japonica subspecies hybrids rice. In this paper, 18 parents and 39 deriving pedigrees were used to reveal the relationship between indica-japonica differentiation and heterosis, and the relationship between heredity diversity and pedigree. The improving effects of yield traits in different breeding stages and genetic effects of yield traits and their interaction by environment were also analyzed. The result would provide scientific foundation for further effective utilization of deriving pedigree of indica-japonica subspecies hybrids rice and hybrid rice combination. The main experimental results are as follows.
1. Differentiation of indica-japonica rice subspecies hybrids deriving pedigree, including 18 parents, 39 deriving pedigrees and 2 control varieties, were determinated by using 58 ILP markers (Intron Length Polymorphism Markers in rice) and Cheng’s Index method. The discrimination value Di and component index of Indica-japonica were selected as qualitative index to distinguish the amplified fragment polymorphism and classify types of indica-japonica. The results show that indica-japonica differentiation existed at the detection sites in the genomic DNA from materials. no indica type (or indica type) was detected with 100% component index. Of 18 parents, 4 japonica, 5 japonica-clinious, 8 indica-clinous and 1 indica were detected to contain 79.69%, 32.08%, 10.94%, and 6.25% of average component indexes of japonica respectively. In 39 deriving pedigrees of rice subspecies indica-japonica, 1 japonica, 11 japonica-clinious, 20 indica-clinous and 7 indica were detected to have 68.70%, 24.62%, 12.92%, and 4.76% of average component indexes of japonica respectively. The component indexes of Nippornbare and 9311, which belong to japonica and indica, were 95.83% and 4.17% respectively. The indexes of japonica rice cultivar C418 and Minghui 502 were 31.25% and 10.42% respectively, but indica cultivar 9308 and Minghui 63 were 33.33% and 12.50% respectively. Further analysis indicated that cultivar Shanyou 63 hybrid, which were derived from interbred from cultivar Minghui 63, had the largest plantation area in the world the heterosis between rice subspecies indica and japonica. There were also significant correlations between component index and Cheng’s index, and the classification of indica and japonica was similar to the dermination of shape index. Therefore, it’s reasonable to use component and determination index of indica or japonica obtained from ILP markers, in order to determinate the differentiation of indica and japonica. It’s vital to utilize the useful genes from rice subspecies indica-japonica.
2. Studies on genetic diversity and genetic relatives of indica-japonica subspecies rice hybrid deriving pedigrees and the parents was conducted by using 221 SSR and pedigree analysis. The results showed that the indexes of genetic diversity, such as polymorphic marker number, polymorphic marker percentage (PPB), effective number of alleles (Ne), observed number of alleles (Na), Shannon diversity index (H) et al., from parents were higher than those from deriving pedigrees, the forth generation deriving pedigree was higher than the second pedigree. And the order of indica and japonica differentiation from high to low is japonica-clinious, indica-clinious, japonica, indica. The deriving pedigrees and parents were divided into 5 clusters by SSR markers, and then the third cluster divided into 4 sub-clusters. There was a significant difference between different indica-japonica types of deriving pedigrees or parents. The cultivars, which were closely relative to cultivar Minghui 63, were gathered at the second sub-cluster, and the third cluster. It’s suggested that most deriving pedigrees could significantly improve the genetic basis of three line restorer. The results of population genetic structure indicates that with the improvement of deriving pedigrees, genetic component was increased and the component ratio of indica or japonica deriving pedigrees tends to be reasonable. Pedigree allele method was used to analyze the transfer of genetic materials from parents in deriving generation. The results showed that the genetic contribution rations of ancestors, such as cultivar Ketan Nangka, Jiananjin, 02428 and Minghui 72, to the first deriving generation were 15.09%, 21.32%, 22.78% and 34.36% respectively, the second generation were 6.05%, 7.09%, 7.98% and 13.41%, respectively, for the third were 3.51%, 4.04%, 4.56% and 7.92% respectively, for the forth were 3.31%, 3.23%, 3.68% and 7.55% respectively. The order of genetic contribution rations of ancestor parents to deriving generations from large to small is cultivar Minghui 72, 02428, cultivar Jiananjin, Ketan Nangka. Each parent including the specific genetic loci was also passed to the deriving offspings, and the genetic contribution to deriving pedigrees decreased with the increase of the deriving generations. Cultivar 92gk729 kept 10 SSR specific loci from ancestor parents after genetic reconstruction, the 5 loci of which could be stably passed to deriving generations with 23.21%, 35.71%, 82.14%, 64.29% and 41.07%. The forth deriving generation inherited 2.21 loci from parents on average. The cluster results of cultivar Ketan Nangka, Jiananjin and 02428 were similar. Parents and their deriving pedigrees, but had a significant difference from cultivar Minghui 72. The results suggested that the different genetic background of deriving pedigrees and helped to understand the genetic structure differentiation of deriving pedigrees.
3. Using 4 indica or japonica sterile line and 54 deriving pedigrees and their parents, 216 cross combinations were generated with incomplete diallel cross design (NC-II), so that the relation between genetic differences of biparental indica-japonica and heterosis of yield characters could be discussed. 96 cross combinations, from 8 three-line and tow-line CMS and 12 rice subspecies differentiation with significant differences, were selected in order to research on the application sphere of genetic differences of parental incica-japonica. 41 ILP markers were used to detect the indica-japonica differentiation. The results revealed that the proportion index of japonica of CMS lines cultivar Chuenjiang 12A, Guangkang 13A, K17A, Zhong 9A, Jin 23A and II-32A was 92.50% (japonica), 9.76% (indica-clinious), 9.76% (indica-clinious), 5% (indica), 7.32% (indica), 7.69% (indica), respectively. The proportion index of japonica of TGMS lines cultivar Peiai 64S, Guangzhan 63S, and SE21S were 24.39 % ( japonica-clinious), 12.50 % ( indica-clinious) and 7.32 % (indica), respectively. Effects of genetic differences of parental indica-japonica on the heterosis of yield traits were diverse for the different materials, but had same trends in different places. Genetic differences of parental indica-japonica had considerable influence on the transgressive heterosis of setting grains per panicle and setting grain percentage, but little influence on the effective panicles and 1000-grain weight. The result of research on the application sphere of genetic differences of parental incica-japonica showed that the character of grain weight per plant was the lowest negative heterosis when the difference of restorer to sterile lines was -5%. When the difference from -5% to 0, the positive heterosis increased, When the difference from 0 to 20%, the positive heterosis was much more than 20%, but the positive heterosis decreased from 20% to 25%. The hybrid rice with strong heterosis could be screened from the restorer lines, which were indica-clinious and japonica-clinious type, when the difference from 0 to 10% and 10% to 20%, respectively. The application range of genetic difference of the parental indica-japonica in two-line hybrid rice was much wider than in three-line hybrid rice. It could be easy to select high yield combinations from the difference of 7.7% to 15%.
4. On the basis of application range, the effects of genetic difference from parental indica-japonica on the heterosis of stalk and photosynthetic characters were further analyzed. The results indicated that there was a significantly or extremely negative correlation between the component index differences of parents and lodging index of heterosis, but no significant correlation between bending moment and stem ellipse section area of heterosis in all cross combinations. There was a significant correlation between the component index differences of parents with breaking resistance and stem wall section area of transgressive heterosis. The component index of parents was significantly correlated with the photosynthetic characters at full heading stage of tansgreesive or mid-parent heterosis, but at maturity there was a positive correlation.
5. By using 8 hybrid lines of subspecies rice and 8 three-line and tow-line sterile lines, two generations of parent and F1 hybrid were selected by using incomplete diallel cross design (NC-Ⅱ), so that the variation trends of phenotypic value of 9 yield characters, genetic effects and heterosis in different deriving pedigrees was analyzed with additive-dominance genetic model. The results of phenotypic value showed that grain weight per plant increased along with the increase of deriving generations. And the change of the second and the forth deriving generation was not significant, but both of them increased much more than the first deriving generation. The total seed per panicles and the setting seeds per panicles of parents and F1 aggrandized, but plant height in the second deriving pedigrees and the third deriving pedigrees changed little, but increased much more than the first deriving generation. The small change of heading stage between deriving generations indicates that transgressive heterosis has good improving effect on controlling growth period of F1. Genetic effect analysis reckoned that additive variance of total seeds per panicles and setting seeds per panicles increased of phenotypic variance, but dominant variance decreased, and the environmental effect was not obvious. The additive variance of growth period and 1000-seeds weight of phenotypic variance in the forth deriving generation was larger than those in the first and second deriving generation, and dominant variance of phenotypic variance was low. The results of heterosis analysis showed that growth period of population average and transgressive heterosis changed less in different deriving generations, and most were negative heterosis. The total seeds per panicles, setting seeds per panicles and plant height decreased, and most were positive heterosis.
6. By using 8 hybrid strains of the forth deriving generation and 8 three-line and tow-line sterile lines, two generations of parent and F1 hybrid were selected with incomplete diallel cross design (NC-Ⅱ), so that the genetic effects and heterosis of yield characters under different types of environments was analyzed by using additive-dominance genetic model, and the value in use of hybrid strains was estimated. The results show that 8 characters were controlled by the main genetic effect, and the main genetic effect variance was from 54.9% to 91.3% of genetic variance except for grain weight per plant. The main genetic effect variance of characters, such as heading stage, effective panicles per plant, total seeds per panicles, setting seeds per panicles and 1000-seeds weight, mainly was the additive variance, whose main genetic effects were 76.8%, 70.9%, 94.3%, 84.95 and 85.8%, respectively. The characters of them could be stably inherited by selection. The main genetic effect variance of characters, such as grain weight per plant, setting percentage, plant height and panicle length, mainly was the dominant variance, which was 68.8%, 78.8%, 56.3% and 61.3% of main genetic effect, respectively. These 4 characters could be utilized by heterosis. The results of heterosis analysis showed that there were significant or extremely significant positive heterosis in 7 characters, except effective panicles per plant and heading stage. The grain weight per plant was the biggest (18.5%), and 1000-grain weight was the smallest, only 3.2%. The stability of population average heterosis of effective panicles, 1000-grain weight and plant height was better than the other 6 characters. There were significant negative population transgressive heterosis in the 6 characters, except for grain weight per plant, plant height and panicle length. The stability of population trangressive heterosis of plant height, heading stage and effective panicle per plant was better than other 6 characters. Analysis of breeding value for breed strains indicated that cultivar Minghui 413 could increase effect for the total seeds per panicles, panicles length and grain weight per plant in hybrid progeny, cultivar Minghui 509 could increase effect for the total seeds per panicles, seeds setting rate, panicles length and grain weight per plant, cultivar Minghui 417 could increase effect for setting percentage, 1000-grain weight, panicles length and grain weight per plant, cultivar Minghui 118 could increase effect for setting percentage and grain weight per plant, cultivar Minghui 502 could increase effect for panicles length, total seeds per panicles and seeds setting rate, cultivar Minghui 503 and 512 could increase effect for total seeds per panicles, Minghui 398 could decrease effect for plant height and increase effect for 1000- grain weight.
引文
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