长江中下游杂交粳稻亲本产量和品质性状优异配合力标记基因型筛选与配合力改良研究
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摘要
全球一半以上的人口以稻米为主食。在耕地面积不断减少、水资源日益匮乏的大环境下,要满足不断增加的世界人口对稻米的需求,只有提高单产。过去30多年的实践已经证明,杂种水稻是提高单产的成功的技术。中国每年种植1700万公顷杂交水稻,占水稻种植总面积的50%左右。杂交籼稻种植面积已占籼稻种植总面积的70%左右。杂交粳稻种植面积仅占粳稻种植总面积840万公顷的3%左右。杂交粳稻还有很大的发展空间。限制杂交粳稻发展速度最主要的因子是竞争优势不够强。提高杂交粳稻竞争优势的关键在于改良恢复系的配合力。本研究在前期工作的基础上,开展了以下4个方面的研究工作：一是利用粳稻6个BT型雄性不育系和12个恢复系按NC Ⅱ遗传交配设计组配72个F1组合,调查8个产量相关性状表型值,分析18个亲本8个产量相关性状的一般配合力效应和特殊配合力效应方差,结合18个亲本115对SSR引物扩增的DNA分子标记数据,筛选18个亲本的单株日产量、单株有效穗数、每穗总粒数、每穗实粒数、千粒重、一次枝梗数、二次枝梗数和穗长等8个产量相关性状优异配合力的SSR标记基因型。二是调查上述72个F1组合的谷粒长、谷粒宽、糙米率、精米率、整精米率、垩白米粒率、垩白度、糊化温度、胶稠度和直链淀粉含量等10个米质性状表型值,分析18个亲本10个米质性状的一般配合力效应和特殊配合力效应方差,结合亲本SSR分子标记数据,筛选18个亲本10个米质性状优异配合力的标记基因型。三是通过调整不育系和恢复系材料类型和数量,扩大亲本群体至不育系7个,恢复系13个,利用同一套SSR引物,扩增这20个亲本SSR分子标记基因型,结合91个组合F1产量性状表型数据,再次鉴定亲本产量性状优异配合力的标记基因型；并与上述第1项研究结果比较,分析亲本群体扩大后各性状优异配合力标记基因型的变化情况；以及2年同一套亲本同一套标记产量相关性状配合力的标记基因型的异同。四是利用上述第1项和第2项研究筛选出的产量和品质相关性状优异配合力标记基因型,通过杂交、回交和SSR分子标记辅助多代选择技术,对江苏省审定组合86优8号的父本恢复系宁恢8号的单株有效穗数和胶稠度性状的配合力、以及优质恢复系157TR-68的单株日均产量配合力进行实际改良和效果评价。获得的主要结果如下。
一、在6个不育系与12个恢复系所配的72个F1组合中,共筛选出31个SSR标记基因型与亲本8个产量相关性状配合力显著相关。其中2个标记基因型同时与亲本6个性状配合力相关；有2个标记基因型同时与亲本5个性状配合力相关；有4个标记基因型同时与亲本4个性状配合力相关；有5个标记基因型同时与亲本3个性状配合力相关；有3个标记基因型同时与亲本2个性状配合力相关；有15个标记基因型与亲本单个性状配合力相关。与亲本4个性状配合力相关的4个标记基因型中,有1个标记基因型RM23～150/160对4个性状的配合力效应都是正的,可使F1的每穗总粒数、单株日产量、穗长和二次枝梗数4个性状值分别增加11.6%、11.2%、10.1%、15.0%。与亲本3个性状配合力相关的5个标记基因型中,有3个标记基因型对亲本的配合力效应是正值。
二、在6个不育系与12个恢复系配制的72个F1组合中,共鉴定出30个SSR标记基因型与亲本10个米质性状配合力显著相关。其中25个与亲本米质性状不良配合力相关,5个与优异配合力相关。标记基因型RM263-175/180和RM444-230/240可以使F1整精米率分别提高3.2%和2.5%。RM3-120/150可以使F1谷粒长缩短2.4%,RM444-180/240可以使F1谷粒宽增加2.1%。RM428-273/294可以使F1植株上的杂交稻米直链淀粉含量减少7.0%。有8个标记基因型同时也影响产量性状配合力。RM3-120/150同时可以使F1的每穗总粒数和每穗实粒数分别增加15.9%和10.9%。RM1211-150/160可使F1的糙米率和精米率分别减少0.9%和1.1%,同时使F1的每穗总粒数和每穗实粒数分别增加21.8.%和20.3%。RM23-150/160可使F1的垩白米粒率和垩白度分别增加44.1%和45.7%,同时使F1的单株日产量和每穗总粒数分别增加11.2%和11.6%。这些结果可用于指导亲本米质性状和产量性状配合力的分子标记辅助改良以及未来杂交粳稻组合配置中的亲本选配。
三、在7个不育系与13个恢复系配制的91个F1组合中,共发现66个SSR标记基因型与亲本产量相关性状配合力显著相关。其中22个与亲本单个性状配合力相关；19个同时与亲本2个性状配合力相关；8个同时与亲本3个性状配合力相关；10个同时与亲本4个性状配合力相关；3个同时与亲本5个性状配合力相关；4个同时与亲本6个性状配合力相关。比较2年同一套亲本同一套标记产量相关性状配合力的标记基因型,发现2年都与同一性状配合力显著相关的标记基因型26个。其中4个与单株有效穗数、8个与每穗总粒数、11个与穗长、3个与每穗实粒数的配合力显著相关。
四、运用优异配合力标记基因型RM208-175/180和RM5556-96/96信息,以武育粳3号R为RM208-180bp和RM5556-96bp条带的供体亲本,改良粳稻恢复系宁恢8号单株有效穗数和精米胶稠度配合力；利用优异配合力标记基因型RM152-165/170信息,以宁恢8号为RM152-170bp条带的供体亲本,改良优质粳稻恢复系宁恢157单株日产量配合力。通过杂交、回交、自交和测交,获得了RM208位点为180bp纯合条带的宁恢8号遗传背景的新恢复系12个。其中8010-4-24、8010-6-5和8010-6-23与对应不育系武3A配组,F1单株有效穗数比对照宁恢8号与武3A所配组合F1平均提高了45%；8010-4-10、8010-4-14和8012-2-9与不育系863A配组,F1单株有效穗数比对照宁恢8号与863A所配组合F1分别增加了65.5%、56.9%和86.2%。获得了RM5556位点为96bp纯合条带的宁恢8号遗传背景的新恢复系6个。其中8009-14-3、8009-14-18、8009-14-29、8013-22-10和8013-22-24与9522A配组,F1植株上所收稻谷的精米胶稠度均显著低于宁恢8号与9522A配组。获得了RM152位点为170bp纯合条带的宁恢157遗传背景的新恢复系8个,其中8014-26-1、8014-26-12、8014-26-21、8014-26-24、8015-7-4和8015-7-17与六千辛A配组,F1单株日产量显著高于对照宁恢157与六千辛A配组,6个F1的单株日产量分别比对照六优157(六千辛-宁恢157F1)增加86.81%、43.06%、78.47%、55.56%、90、97%和82.64%。
综合上述亲本配合力标记基因型筛选结果,发现与产量相关性状配合力相关的SSR标记基因型数目较多,三分之一标记基因型与2个及以上性状的配合力同时相关。那些与改良目标一致的多性状配合力相关位点可能有利于亲本多个性状配合力同时进行改良。年份间重复检测到的与同一性状配合力显著相关的标记基因型,对F1性状值的效应是同向的,增量率是相近的。这些年份间表现稳定的优异配合力标记基因型用于粳稻恢复系配合力改良预计可以收到期望的效果。与亲本米质性状配合力相关的SSR标记基因型中,80%的位点处于杂合状态时,对F1植株上所收稻谷的精米品质有不利的影响,这可能与F1植株上所收籽粒基因型发生分离有关。因此在影响米质性状的位点上,双亲应当具有相同的等位基因。利用优异配合力标记基因型信息,通过染色体等位片段置换,改良亲本产量相关性状配合力的效果大于改良亲本米质性状配合力的效果。武3A/8010-4-24和863A/8012-2-9是单株有效穗数、单株日产量和单株产量增加最多且株高生育期与对照差异不显著的两个组合,有望在生产上应用。
Rice (Oryza sativa L.) is a staple food for more than half of the world population. The increasing demand for rice in the world caused by population increase can only be met by enhancing yield per unit area under the environment that both arable land acrage and water resources are decreasing. Hybrid rice is a proven and successful technology for rice production over the past three decades. Seventeen million hectares of hybrid rice are grown in China, occupying about50%of total rice area planted annually. Hybrid indica rice accounts for70%of total indica rice area. The total area of japonica rice grown annually in China is8.4million hectares, and only3%is occupied by japonica hybrid rice. Japonica hybrids still have great potential to development. The major reason for this situation was that competitive heterosis of hybrid cultivar was not conspicuous in yield and quality, compared with conventional cultivar in japonica rice. The key of enhancing competitive heterosis of hybrid cultivar in japonica rice was to improve combining ability (CA) of yield related traits and quality traits of restorer lines. On the basis of previous studies, four aspects of research are carried out in this study. Firstly,72F1combinations were made from6cytoplasmic male sterile (CMS) lines and12restorer lines according to North Carolina Genetic Mating Design Ⅱ(NCⅡ). Eight yield related traits of the72F1combinations were investigated, and general combining ability (GCA) effects and variance of special combining ability (VSCA) of the8traits of the18parents was analyzed. SSR marker genotypes for elite CA of the eight traits, i. e., daily yield per plant (DYP), panicles per plant (PP), total spikelets per panicle (TSP), filled spikelets per panicle (FSP) and1000-grain weight (TGW), primary branch number per panicle(PBN), second branch number per panicle(SBN) and panicle length(PL), were screened by combining the SSR genotyping data with CA data of the18parents. Secondly,10quality traits, namely, brown rice rate, milled rice rate, head rice rate, unhulled grain length, grain width, percentage of chalky grain, chalkiness degree, gelatinization temperature, gel consistency, and amylose contents, of the72F1S made from the same6CMS lines and12restorer lines as above were measured. And the CA of the10quality traits of the18parents was analyzed. SSR marker genotypes for elite CA of the10quality traits were screened by the same method as mentioned above. Thirdly, SSR marker genotypes for elite CA of the grain yield related traits were re-identified by using phenotype values of91F1combinations made from7CMS lines and13restorer lines and genotype values of the20parents. The115SSR primers used for genotyping the20parents are the same as above. Among the7CMS lines and13restorer lines used in this part,6CMS lines and10restorer lines were identical to those used in previous report and in part1and2of this study. SSR marker genotypes for elite CA of the traits were compared with that in enlarged parent population. And consistency of SSR marker genotypes for elite CA of the grain yield related traits across years was analyzed using the same set of parents and the same set of SSR primers. Forthly, improvement of CA for productive panicles per plant and gel consistency in a restorer line Ninghui8and for daily average grain yield in another restorer line157TR-68were carried out by using hybridization, backcrossing and assisted selection of SSR markers identified in this study. The main results obtained were the follows:
1. Thirty-one SSR marker genotypes were significantly associated with CA of the eight traits, using the dataset of72F1s made from the six CMS lines and twelve restorer lines. Out of the thirty-one, fifteen were associated with one trait, three for two traits, five for three traits, four for four traits, two for five traits, and two for six traits. Among the four marker genotypes associated with four traits simultaneously, RM23-150/160showed positive effects of CA in all the four traits, increasing11.6%of total spikelets per panicle,11.2%of daily yield per plant,10.1%of panicle length and15.0%of second branch number per panicle in F1, respectively. Among the five marker genotypes associated with three traits simultaneously, three showed positive effects of CA for all the three traits.
2. A total of30SSR marker genotypes were significantly associated with CA of the10quality traits, namely, brown rice rate, milled rice rate, head rice rate, grain length, grain width, percentage of chalky grain, chalkiness degree, gelatinization temperature, gel consistency, and amylose contents, using the dataset of72F1s made from the same6CMS lines and12restorer lines as above. Twenty-five of them were marker genotypes for unfavorable CA of quality traits. The remaining five marker genotypes were associated with elite CA for quality traits of parents. RM263-175/180and RM444-230/240were marker genotypes of elite CA for head rice rate, which increased3.2%and2.5%of head rice rate in F1, respectively. RM3-120/150shortened grain length value in F1by2.4%. RM444-180/240increased grain width trait value of F1by2.1%. RM428-273/294decreased amylose content trait value of hybrid rice harvested from F1plants by7.0%. Among the SSR marker genotypes associated with CA of the10quality traits, eight marker genotypes also affected CA of yield components simultaneously. RM3-120/150increased simultaneously total spikelets per plant and filled spikelets per plant in F1by15.9%and10.9%, respectively. RM1211-150/160decreased trait values of brown rice rate and milled rice rate in F1by0.9%and1.1%, respectively, and simultaneously increased trait values of total spikelets per plant and filled spikelets per plant in F1by21.8%and20.4%. RM23-150/160increased trait values of percentage of chalky grain and chalkiness degree in F1by44.1%and45.7%, and simultaneously increased daily yield per plant and total spikelets per plant in F1by11.2%and11.6%, respectively.
3. Sixty-six SSR marker genotypes were significantly associated with CA of9yield related traits, namely, DYP, PP, TSP, FSP, TGW, days to heanding (DTH), plant height (PH), PL and grains per cm panicle (GPCP), using the dataset of91F1s derived from7BT-type japonica male sterile lines and13restorer lines in rice made with NCII design. Among the66SSR marker genotypes detected, Twenty-two were significantly associated for only one trait, nineteen for two traits, eight for three traits, ten for four traits, three for five traits and four for six traits. Comparing the marker genotypes identified from the identical16parents and115SSR markers used in2009and in2010,26marker genotypes were found associated significantly with the same trait in both years. Among them, four for PP, eight for TSP, eleven for PL, three for FSP.
4. By hybridizing, bachcrossing, selfing and testcrossing.12new restorers with180bp homozygous at marker locus RM208in background of Ninghui8were obtained. Among them, combining ability for productive panicle per plant in8010-4-24,8010-6-5and8010-6-23increased by45%averagely, compared with that of check restorer Ninghui8, when crossed with CMS line Wu3A. And CA for productive panicle per plant in8010-4-10,8010-4-14and8012-2-9enhanced by65.5%,56.9%and86.2%, respectively, compared with that of Ninghui8, when crossed with CMS line863A. Six new restorers with96bp homozygous at marker locus RM5556in background of Ninghui8were obtained. CA for gel consistency of milled rice in8009-14-3,8009-14-18,8009-14-29,8013-22-10and8013-22-24was significantly lower than that of check restorer Ninghui8, when crossed with CMS line9522A. Eight new restorers with170bp homozygous at marker locus RM152in background of Ninghui157were bred. CA for daily grain yield per plant in8014-26-1,8014-26-12,8014-26-21,8014-26-24,8015-7-4and8015-7-17was significantly higher than that of check restorer Ninghui157, when crossed with CMS line Liuqianxin A. Daily grain yield per plant of the6F1increased by86.8%,43.1%,78.5%,55.6%,91.0%and82.6%, compared with check Liuyou157.
Summaring the screening results, we found many SSR loci associated with CA of yield related traits in parents, and about33%of the loci were associated with two or more than two traits. Those loci associated with CA of multi-traits consistent with breeding objectives might facilitate CA improvement for multi-traits simultaneously. Marker genotypes that were repeatly detected across years and were associated significantly with CA of the same trait showed the same effect to F1trait value in direction, and the percentage of increment was almost the same in different years. Expect results may be obtained by using the stable marker genotypes across years for improving CA of restorer lines in japonica rice. Among the SSR marker genotype loci associated with CA of quality traits,80%of the heterozygous loci were unfavorable to quality of milled rice obtained from the F1plants. This phenomenon may be related to the genotype segregation of grains harvested from from F1plants. Thus two parents with identical alleles at loci affecting quality traits are desirable. The efficiency of improving CA of yield related traits was better than that of quality traits by allele substitution according to information of elite CA marker genetyps. Wu3A/8010-4-24and863A/8012-2-9had more productive panicles per plant and higher daily average grain yield than their corresponding check cultivars, and there was no significant difference between the combinations and their checks in plant height and growth duration. These two combinations, therefore, are hopeful to be used in production.
一、在6个不育系与12个恢复系所配的72个F1组合中,共筛选出31个SSR标记基因型与亲本8个产量相关性状配合力显著相关。其中2个标记基因型同时与亲本6个性状配合力相关；有2个标记基因型同时与亲本5个性状配合力相关；有4个标记基因型同时与亲本4个性状配合力相关；有5个标记基因型同时与亲本3个性状配合力相关；有3个标记基因型同时与亲本2个性状配合力相关；有15个标记基因型与亲本单个性状配合力相关。与亲本4个性状配合力相关的4个标记基因型中,有1个标记基因型RM23～150/160对4个性状的配合力效应都是正的,可使F1的每穗总粒数、单株日产量、穗长和二次枝梗数4个性状值分别增加11.6%、11.2%、10.1%、15.0%。与亲本3个性状配合力相关的5个标记基因型中,有3个标记基因型对亲本的配合力效应是正值。
二、在6个不育系与12个恢复系配制的72个F1组合中,共鉴定出30个SSR标记基因型与亲本10个米质性状配合力显著相关。其中25个与亲本米质性状不良配合力相关,5个与优异配合力相关。标记基因型RM263-175/180和RM444-230/240可以使F1整精米率分别提高3.2%和2.5%。RM3-120/150可以使F1谷粒长缩短2.4%,RM444-180/240可以使F1谷粒宽增加2.1%。RM428-273/294可以使F1植株上的杂交稻米直链淀粉含量减少7.0%。有8个标记基因型同时也影响产量性状配合力。RM3-120/150同时可以使F1的每穗总粒数和每穗实粒数分别增加15.9%和10.9%。RM1211-150/160可使F1的糙米率和精米率分别减少0.9%和1.1%,同时使F1的每穗总粒数和每穗实粒数分别增加21.8.%和20.3%。RM23-150/160可使F1的垩白米粒率和垩白度分别增加44.1%和45.7%,同时使F1的单株日产量和每穗总粒数分别增加11.2%和11.6%。这些结果可用于指导亲本米质性状和产量性状配合力的分子标记辅助改良以及未来杂交粳稻组合配置中的亲本选配。
三、在7个不育系与13个恢复系配制的91个F1组合中,共发现66个SSR标记基因型与亲本产量相关性状配合力显著相关。其中22个与亲本单个性状配合力相关；19个同时与亲本2个性状配合力相关；8个同时与亲本3个性状配合力相关；10个同时与亲本4个性状配合力相关；3个同时与亲本5个性状配合力相关；4个同时与亲本6个性状配合力相关。比较2年同一套亲本同一套标记产量相关性状配合力的标记基因型,发现2年都与同一性状配合力显著相关的标记基因型26个。其中4个与单株有效穗数、8个与每穗总粒数、11个与穗长、3个与每穗实粒数的配合力显著相关。
四、运用优异配合力标记基因型RM208-175/180和RM5556-96/96信息,以武育粳3号R为RM208-180bp和RM5556-96bp条带的供体亲本,改良粳稻恢复系宁恢8号单株有效穗数和精米胶稠度配合力；利用优异配合力标记基因型RM152-165/170信息,以宁恢8号为RM152-170bp条带的供体亲本,改良优质粳稻恢复系宁恢157单株日产量配合力。通过杂交、回交、自交和测交,获得了RM208位点为180bp纯合条带的宁恢8号遗传背景的新恢复系12个。其中8010-4-24、8010-6-5和8010-6-23与对应不育系武3A配组,F1单株有效穗数比对照宁恢8号与武3A所配组合F1平均提高了45%；8010-4-10、8010-4-14和8012-2-9与不育系863A配组,F1单株有效穗数比对照宁恢8号与863A所配组合F1分别增加了65.5%、56.9%和86.2%。获得了RM5556位点为96bp纯合条带的宁恢8号遗传背景的新恢复系6个。其中8009-14-3、8009-14-18、8009-14-29、8013-22-10和8013-22-24与9522A配组,F1植株上所收稻谷的精米胶稠度均显著低于宁恢8号与9522A配组。获得了RM152位点为170bp纯合条带的宁恢157遗传背景的新恢复系8个,其中8014-26-1、8014-26-12、8014-26-21、8014-26-24、8015-7-4和8015-7-17与六千辛A配组,F1单株日产量显著高于对照宁恢157与六千辛A配组,6个F1的单株日产量分别比对照六优157(六千辛-宁恢157F1)增加86.81%、43.06%、78.47%、55.56%、90、97%和82.64%。
综合上述亲本配合力标记基因型筛选结果,发现与产量相关性状配合力相关的SSR标记基因型数目较多,三分之一标记基因型与2个及以上性状的配合力同时相关。那些与改良目标一致的多性状配合力相关位点可能有利于亲本多个性状配合力同时进行改良。年份间重复检测到的与同一性状配合力显著相关的标记基因型,对F1性状值的效应是同向的,增量率是相近的。这些年份间表现稳定的优异配合力标记基因型用于粳稻恢复系配合力改良预计可以收到期望的效果。与亲本米质性状配合力相关的SSR标记基因型中,80%的位点处于杂合状态时,对F1植株上所收稻谷的精米品质有不利的影响,这可能与F1植株上所收籽粒基因型发生分离有关。因此在影响米质性状的位点上,双亲应当具有相同的等位基因。利用优异配合力标记基因型信息,通过染色体等位片段置换,改良亲本产量相关性状配合力的效果大于改良亲本米质性状配合力的效果。武3A/8010-4-24和863A/8012-2-9是单株有效穗数、单株日产量和单株产量增加最多且株高生育期与对照差异不显著的两个组合,有望在生产上应用。
Rice (Oryza sativa L.) is a staple food for more than half of the world population. The increasing demand for rice in the world caused by population increase can only be met by enhancing yield per unit area under the environment that both arable land acrage and water resources are decreasing. Hybrid rice is a proven and successful technology for rice production over the past three decades. Seventeen million hectares of hybrid rice are grown in China, occupying about50%of total rice area planted annually. Hybrid indica rice accounts for70%of total indica rice area. The total area of japonica rice grown annually in China is8.4million hectares, and only3%is occupied by japonica hybrid rice. Japonica hybrids still have great potential to development. The major reason for this situation was that competitive heterosis of hybrid cultivar was not conspicuous in yield and quality, compared with conventional cultivar in japonica rice. The key of enhancing competitive heterosis of hybrid cultivar in japonica rice was to improve combining ability (CA) of yield related traits and quality traits of restorer lines. On the basis of previous studies, four aspects of research are carried out in this study. Firstly,72F1combinations were made from6cytoplasmic male sterile (CMS) lines and12restorer lines according to North Carolina Genetic Mating Design Ⅱ(NCⅡ). Eight yield related traits of the72F1combinations were investigated, and general combining ability (GCA) effects and variance of special combining ability (VSCA) of the8traits of the18parents was analyzed. SSR marker genotypes for elite CA of the eight traits, i. e., daily yield per plant (DYP), panicles per plant (PP), total spikelets per panicle (TSP), filled spikelets per panicle (FSP) and1000-grain weight (TGW), primary branch number per panicle(PBN), second branch number per panicle(SBN) and panicle length(PL), were screened by combining the SSR genotyping data with CA data of the18parents. Secondly,10quality traits, namely, brown rice rate, milled rice rate, head rice rate, unhulled grain length, grain width, percentage of chalky grain, chalkiness degree, gelatinization temperature, gel consistency, and amylose contents, of the72F1S made from the same6CMS lines and12restorer lines as above were measured. And the CA of the10quality traits of the18parents was analyzed. SSR marker genotypes for elite CA of the10quality traits were screened by the same method as mentioned above. Thirdly, SSR marker genotypes for elite CA of the grain yield related traits were re-identified by using phenotype values of91F1combinations made from7CMS lines and13restorer lines and genotype values of the20parents. The115SSR primers used for genotyping the20parents are the same as above. Among the7CMS lines and13restorer lines used in this part,6CMS lines and10restorer lines were identical to those used in previous report and in part1and2of this study. SSR marker genotypes for elite CA of the traits were compared with that in enlarged parent population. And consistency of SSR marker genotypes for elite CA of the grain yield related traits across years was analyzed using the same set of parents and the same set of SSR primers. Forthly, improvement of CA for productive panicles per plant and gel consistency in a restorer line Ninghui8and for daily average grain yield in another restorer line157TR-68were carried out by using hybridization, backcrossing and assisted selection of SSR markers identified in this study. The main results obtained were the follows:
1. Thirty-one SSR marker genotypes were significantly associated with CA of the eight traits, using the dataset of72F1s made from the six CMS lines and twelve restorer lines. Out of the thirty-one, fifteen were associated with one trait, three for two traits, five for three traits, four for four traits, two for five traits, and two for six traits. Among the four marker genotypes associated with four traits simultaneously, RM23-150/160showed positive effects of CA in all the four traits, increasing11.6%of total spikelets per panicle,11.2%of daily yield per plant,10.1%of panicle length and15.0%of second branch number per panicle in F1, respectively. Among the five marker genotypes associated with three traits simultaneously, three showed positive effects of CA for all the three traits.
2. A total of30SSR marker genotypes were significantly associated with CA of the10quality traits, namely, brown rice rate, milled rice rate, head rice rate, grain length, grain width, percentage of chalky grain, chalkiness degree, gelatinization temperature, gel consistency, and amylose contents, using the dataset of72F1s made from the same6CMS lines and12restorer lines as above. Twenty-five of them were marker genotypes for unfavorable CA of quality traits. The remaining five marker genotypes were associated with elite CA for quality traits of parents. RM263-175/180and RM444-230/240were marker genotypes of elite CA for head rice rate, which increased3.2%and2.5%of head rice rate in F1, respectively. RM3-120/150shortened grain length value in F1by2.4%. RM444-180/240increased grain width trait value of F1by2.1%. RM428-273/294decreased amylose content trait value of hybrid rice harvested from F1plants by7.0%. Among the SSR marker genotypes associated with CA of the10quality traits, eight marker genotypes also affected CA of yield components simultaneously. RM3-120/150increased simultaneously total spikelets per plant and filled spikelets per plant in F1by15.9%and10.9%, respectively. RM1211-150/160decreased trait values of brown rice rate and milled rice rate in F1by0.9%and1.1%, respectively, and simultaneously increased trait values of total spikelets per plant and filled spikelets per plant in F1by21.8%and20.4%. RM23-150/160increased trait values of percentage of chalky grain and chalkiness degree in F1by44.1%and45.7%, and simultaneously increased daily yield per plant and total spikelets per plant in F1by11.2%and11.6%, respectively.
3. Sixty-six SSR marker genotypes were significantly associated with CA of9yield related traits, namely, DYP, PP, TSP, FSP, TGW, days to heanding (DTH), plant height (PH), PL and grains per cm panicle (GPCP), using the dataset of91F1s derived from7BT-type japonica male sterile lines and13restorer lines in rice made with NCII design. Among the66SSR marker genotypes detected, Twenty-two were significantly associated for only one trait, nineteen for two traits, eight for three traits, ten for four traits, three for five traits and four for six traits. Comparing the marker genotypes identified from the identical16parents and115SSR markers used in2009and in2010,26marker genotypes were found associated significantly with the same trait in both years. Among them, four for PP, eight for TSP, eleven for PL, three for FSP.
4. By hybridizing, bachcrossing, selfing and testcrossing.12new restorers with180bp homozygous at marker locus RM208in background of Ninghui8were obtained. Among them, combining ability for productive panicle per plant in8010-4-24,8010-6-5and8010-6-23increased by45%averagely, compared with that of check restorer Ninghui8, when crossed with CMS line Wu3A. And CA for productive panicle per plant in8010-4-10,8010-4-14and8012-2-9enhanced by65.5%,56.9%and86.2%, respectively, compared with that of Ninghui8, when crossed with CMS line863A. Six new restorers with96bp homozygous at marker locus RM5556in background of Ninghui8were obtained. CA for gel consistency of milled rice in8009-14-3,8009-14-18,8009-14-29,8013-22-10and8013-22-24was significantly lower than that of check restorer Ninghui8, when crossed with CMS line9522A. Eight new restorers with170bp homozygous at marker locus RM152in background of Ninghui157were bred. CA for daily grain yield per plant in8014-26-1,8014-26-12,8014-26-21,8014-26-24,8015-7-4and8015-7-17was significantly higher than that of check restorer Ninghui157, when crossed with CMS line Liuqianxin A. Daily grain yield per plant of the6F1increased by86.8%,43.1%,78.5%,55.6%,91.0%and82.6%, compared with check Liuyou157.
Summaring the screening results, we found many SSR loci associated with CA of yield related traits in parents, and about33%of the loci were associated with two or more than two traits. Those loci associated with CA of multi-traits consistent with breeding objectives might facilitate CA improvement for multi-traits simultaneously. Marker genotypes that were repeatly detected across years and were associated significantly with CA of the same trait showed the same effect to F1trait value in direction, and the percentage of increment was almost the same in different years. Expect results may be obtained by using the stable marker genotypes across years for improving CA of restorer lines in japonica rice. Among the SSR marker genotype loci associated with CA of quality traits,80%of the heterozygous loci were unfavorable to quality of milled rice obtained from the F1plants. This phenomenon may be related to the genotype segregation of grains harvested from from F1plants. Thus two parents with identical alleles at loci affecting quality traits are desirable. The efficiency of improving CA of yield related traits was better than that of quality traits by allele substitution according to information of elite CA marker genetyps. Wu3A/8010-4-24and863A/8012-2-9had more productive panicles per plant and higher daily average grain yield than their corresponding check cultivars, and there was no significant difference between the combinations and their checks in plant height and growth duration. These two combinations, therefore, are hopeful to be used in production.
引文
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