小粒矮秆水稻在杂交水稻工程化制种中的应用基础研究
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摘要
本文基于对新发现的小粒矮杆材料潇湘矮的认识,提出了杂交水稻工程化制种的设想并开展了相关研究。研究发现潇湘矮的小粒矮生性连锁并受一对株高隐性质量基因rRH控制,该基因位于第5条染色体RM249上游8.4cM距离；通过对潇湘矮的改造,选育出一批光温敏核不育材料,其中171S综合性状表现较好；以171S进行混合直播制种试验,探明了一些制约因子。为实现机械化制种,文章对潇湘矮背景的小粒特性进行研究并设计出了效果良好的籽粒分离机械。主要研究结果如下：
1、潇湘矮特征特性：潇湘矮属迟熟早稻,长沙早季播种,播始历期90d左右,农艺性状类似于直立穗水稻类型,株高55cm,千粒重11.5g左右,穗大粒多,潇湘矮与半矮杆大粒品种杂交F1表现为半矮杆大粒,与高秆大粒杂交F1表现为高秆大粒；用广亲和测验种进行测交,F1结实率均达到75%左右或以上,具有与培矮64相当的广亲和性,潇湘矮籽粒与普通水稻品种籽粒粒重性状差异较大,可以不同粒型混合样本分离开来。
2、潇湘矮小粒矮生性的遗传规律：潇湘矮与半矮杆大粒湘早籼143杂交,F2分离群体半矮杆大粒与矮杆小粒表现为3：1,潇湘矮与高秆大粒材料粒谷早杂交,F2分离出高秆大粒、半矮秆大粒、半矮秆小粒及矮秆小粒四种类型,其比例为9：3：3：1,说明潇湘矮的小粒与矮秆性状属于一因多效或连锁遗传。当小粒半矮杆突变体与半矮杆材料杂交,表现高秆大粒,说明,小粒半矮杆突变体含有高秆抑制基因,由此推断,潇湘矮的小粒矮杆性状是由一对抑制基因在sd条件下的表现,换言之,潇湘矮的小粒矮生性连锁并受一对株高隐性质量基因rRH控制。
3、潇湘矮小粒矮生性的基因定位：为定位rRH基因位点,通过基于BSA的PCR法筛选了水稻12条染色体的部分SSR标记,将来自潇湘矮的半矮秆基因定位到第5条染色体上。经比对测算,其位于RM249上游8.4cM,有别于其下游的sdg。
4、光温敏核不育系选育：用杂交育种手段将光敏核不育特性转入到潇湘矮中,经过多代加压选择,得到了一批光温敏核不育材料,采用回交、与野生稻杂交和离子辐射等方法改良潇湘矮株叶型及提高其柱头外露率。获得了9sy220、9sy507、9d82等一批综合性状较好的不育材料。
5、高产强优势组合选配：以171S为研究对象,进行广泛配组,测定其配合力,结果表明171S配合力较好,可以选配出高产、生育期适当的强优组合。
6、混合直播制种初探：以171S为研究对象,选用生育期与其相近的恢复系440进行混合直播制种,父母本粒数按1：10；1：20；1：30直播,试验按三水平三重复进行。结果表明,171S平均异交结实率为13.71%,三个水平异交结实率差异不显著。文章分析了异交结实率偏低的主要因素：(1)父母本对GA3敏感程度不一致,GA3使用量不易控制；(2)171S剑叶长且宽,影响花粉接受；(3)171S花期不集中。
7、分离机械设计：以潇湘矮为背景的小粒材料进行籽粒分离试验,试验表明,用3.5mm圆孔筛筛选3次以上,可以将小粒与普通大粒(千粒重25g左右)分离,且分离后种子纯度可达到99%以上,符合国家水稻良种纯度指标,为杂交稻混合直播制种、机械混收并分离父母本种子提供可能。依据窝眼型孔的分选原理,利用籽粒长度差异,确定型孔规格,并完善相关设计,制作出分离机械,且分离效果良好。
The assumptions for engineering-oriented seed production of hybrid rice were proposed and the related researches were conducted based on the knowledge of small grain and dwarf indica rice variety-Xiaoxiangai (XXA). The researches results indicated that the small grain and dwarf gene of XXA was linked and controlled by a pair of plant height recessive quality gene-rRH, and this gene located at 8.4 cM upstream from the-RM249 in the fifth chromosome; through improvement of XXA, a passel of photo-thermo-sensitive genic male sterile lines were selected, and the comprehensive characters of 171S were the best among them; using 171S to conduct mixed direct-sowing seed production experiment, and some restrictive factors were clear. To realize mechanized seed production, the small grain character of XXA was researched and the grains separating mechanism with great effect was designed. The main research results were showed as follows:
1. Characters of XXA:XXA is kind of late maturity early rice; it sowing at early season in Changsha, and the duration from seeding to heading was about 90 days; the agronomic traits of XXA were similar to that of erect panicle rice varieties, which plant height was 55 cm,1000-grain weight was around 11.5 g, large panicle and many grains; the hybrid F1 that crossed by XXA and semi-dwarf and large grain variety showed as semi-dwarf and large grain; the hybrid F1that crossed by XXA and high stalk and large grain variety showed as high stalk and large grain; using wide compatibility testing variety to test cross, the setting percentage of F1 reached 75% or more, which wide compatibility was similar to Peiai 64's; the difference between seed weight for XXA and normal rice variety was large, it could been separated grains of XXA from mixed samples.
2. The genetic regularity of small grain and dwarf characters of XXA:when XXA crossed with semi-dwarf and large grain variety-Xiangzaoxian 143, the proportion of semi-dwarf and large grain variety and dwarf and small grain variety that isolated from F2 colony was 3:1; when XXA crossed with high stalk and large grain variety-Liguzao, the proportion of high stalk and large grain variety, semi-dwarf and large grain variety, semi-dwarf and small grain variety and dwarf and small grain variety that isolated from F2 colony was 9:3:3:1; these results indicated that the characters of small grain and dwarf of XXA belonged to pleiotropism or linkage inheritance. When small grain and semi-dwarf mutant crossed with semi-dwarf variety, the progenies showed character of high stalk and large grain, it indicated that small grain and semi-dwarf mutant containing high stalk inhibitory gene. Theses results revealed that the characters of small grain and dwarf of XXA is kind of expression that a pair of inhibitory gene under sd condition, in other words, the characters of small grain and dwarf of XXA were linked and controlled by a pair of plant height recessive quality gene-rRH.
3. Gene localization of small grain and dwarf characters of XXA:In order to locate rRH gene loci, partial SSR markers of 12 rice chromosomes were selected by PCR that based on BSA, and the semi-dwarf gene of XXA was located to the fifth chromosome. Through intercomparison and calculation, the semi-dwarf gene of XXA located at 8.4 cM upstream from- RM249 in the fifth chromosome, which was different from the sdg downriver from the RM249.
4. breeding male sterile line:Using cross breeding method to transport character of photoperiod sensitive genic male sterile into XXA, then through multi-generation pressurized selecting, a passel of photo-thermo-sensitive genic male sterile lines were gotten; and using methods that backcross, hybrid with wild rice and ionizing radiation, etc. to improve shapes of plant and leaf of XXA and increase its stigma exsertion rate. A passel of male sterile lines with great comprehensive characters were obtained, which were 9sy220,9sy507 and 9d82, etc..
5. Quality combinations with characters of high-yield could be selected:Taking 171S as research object match group widely to determine its combining ability, the results showed that 171S has great combining ability, and quality combinations with characters of high-yield and appropriate growth period could be selected.
6. Preliminary Studies on mixed direct-sowing seed production:Taking 171S as research object and taking restorer line 440, which has similar growth period with 171S's, to conduct mixed direct-sowing seed production; direct sowing experiments were conducted with parents'grain proportion of 1:10,1:20 and 1:30 and repeated three times. The results showed that the average outcrossing rate of 171S was 13.71%, and the differences of outcrossing rates among three levels were not significant. Several factors for low outcrossing rate were analyzed as follows:first, the sensitive degrees of GA3 on parents are different, and the applying amount of GA3 is not easy to control; second, the flag leaves of 171S are long and wide, which affecting pollination; third, the florescence of 171S is not concentrated.
7. Separating machine on the design:Taking XXA as small grain sample to conduct separating experiment for grains and seeds. The results showed that using 3.5 mm circular screen to sift over three times, the small grain and normal large grain (1000-grain weight is around 25 g) could be separated, and the purity of separated seeds reached over 99% that accord with national standards of rice commercial seeds. Based on the separating theory of indented mold-hole, the separating machine had been made by utilizing the different lengths between seeds to ensure the characteristics of mold hole and perfecting related designs, and it has great separating effect.
1、潇湘矮特征特性：潇湘矮属迟熟早稻,长沙早季播种,播始历期90d左右,农艺性状类似于直立穗水稻类型,株高55cm,千粒重11.5g左右,穗大粒多,潇湘矮与半矮杆大粒品种杂交F1表现为半矮杆大粒,与高秆大粒杂交F1表现为高秆大粒；用广亲和测验种进行测交,F1结实率均达到75%左右或以上,具有与培矮64相当的广亲和性,潇湘矮籽粒与普通水稻品种籽粒粒重性状差异较大,可以不同粒型混合样本分离开来。
2、潇湘矮小粒矮生性的遗传规律：潇湘矮与半矮杆大粒湘早籼143杂交,F2分离群体半矮杆大粒与矮杆小粒表现为3：1,潇湘矮与高秆大粒材料粒谷早杂交,F2分离出高秆大粒、半矮秆大粒、半矮秆小粒及矮秆小粒四种类型,其比例为9：3：3：1,说明潇湘矮的小粒与矮秆性状属于一因多效或连锁遗传。当小粒半矮杆突变体与半矮杆材料杂交,表现高秆大粒,说明,小粒半矮杆突变体含有高秆抑制基因,由此推断,潇湘矮的小粒矮杆性状是由一对抑制基因在sd条件下的表现,换言之,潇湘矮的小粒矮生性连锁并受一对株高隐性质量基因rRH控制。
3、潇湘矮小粒矮生性的基因定位：为定位rRH基因位点,通过基于BSA的PCR法筛选了水稻12条染色体的部分SSR标记,将来自潇湘矮的半矮秆基因定位到第5条染色体上。经比对测算,其位于RM249上游8.4cM,有别于其下游的sdg。
4、光温敏核不育系选育：用杂交育种手段将光敏核不育特性转入到潇湘矮中,经过多代加压选择,得到了一批光温敏核不育材料,采用回交、与野生稻杂交和离子辐射等方法改良潇湘矮株叶型及提高其柱头外露率。获得了9sy220、9sy507、9d82等一批综合性状较好的不育材料。
5、高产强优势组合选配：以171S为研究对象,进行广泛配组,测定其配合力,结果表明171S配合力较好,可以选配出高产、生育期适当的强优组合。
6、混合直播制种初探：以171S为研究对象,选用生育期与其相近的恢复系440进行混合直播制种,父母本粒数按1：10；1：20；1：30直播,试验按三水平三重复进行。结果表明,171S平均异交结实率为13.71%,三个水平异交结实率差异不显著。文章分析了异交结实率偏低的主要因素：(1)父母本对GA3敏感程度不一致,GA3使用量不易控制；(2)171S剑叶长且宽,影响花粉接受；(3)171S花期不集中。
7、分离机械设计：以潇湘矮为背景的小粒材料进行籽粒分离试验,试验表明,用3.5mm圆孔筛筛选3次以上,可以将小粒与普通大粒(千粒重25g左右)分离,且分离后种子纯度可达到99%以上,符合国家水稻良种纯度指标,为杂交稻混合直播制种、机械混收并分离父母本种子提供可能。依据窝眼型孔的分选原理,利用籽粒长度差异,确定型孔规格,并完善相关设计,制作出分离机械,且分离效果良好。
The assumptions for engineering-oriented seed production of hybrid rice were proposed and the related researches were conducted based on the knowledge of small grain and dwarf indica rice variety-Xiaoxiangai (XXA). The researches results indicated that the small grain and dwarf gene of XXA was linked and controlled by a pair of plant height recessive quality gene-rRH, and this gene located at 8.4 cM upstream from the-RM249 in the fifth chromosome; through improvement of XXA, a passel of photo-thermo-sensitive genic male sterile lines were selected, and the comprehensive characters of 171S were the best among them; using 171S to conduct mixed direct-sowing seed production experiment, and some restrictive factors were clear. To realize mechanized seed production, the small grain character of XXA was researched and the grains separating mechanism with great effect was designed. The main research results were showed as follows:
1. Characters of XXA:XXA is kind of late maturity early rice; it sowing at early season in Changsha, and the duration from seeding to heading was about 90 days; the agronomic traits of XXA were similar to that of erect panicle rice varieties, which plant height was 55 cm,1000-grain weight was around 11.5 g, large panicle and many grains; the hybrid F1 that crossed by XXA and semi-dwarf and large grain variety showed as semi-dwarf and large grain; the hybrid F1that crossed by XXA and high stalk and large grain variety showed as high stalk and large grain; using wide compatibility testing variety to test cross, the setting percentage of F1 reached 75% or more, which wide compatibility was similar to Peiai 64's; the difference between seed weight for XXA and normal rice variety was large, it could been separated grains of XXA from mixed samples.
2. The genetic regularity of small grain and dwarf characters of XXA:when XXA crossed with semi-dwarf and large grain variety-Xiangzaoxian 143, the proportion of semi-dwarf and large grain variety and dwarf and small grain variety that isolated from F2 colony was 3:1; when XXA crossed with high stalk and large grain variety-Liguzao, the proportion of high stalk and large grain variety, semi-dwarf and large grain variety, semi-dwarf and small grain variety and dwarf and small grain variety that isolated from F2 colony was 9:3:3:1; these results indicated that the characters of small grain and dwarf of XXA belonged to pleiotropism or linkage inheritance. When small grain and semi-dwarf mutant crossed with semi-dwarf variety, the progenies showed character of high stalk and large grain, it indicated that small grain and semi-dwarf mutant containing high stalk inhibitory gene. Theses results revealed that the characters of small grain and dwarf of XXA is kind of expression that a pair of inhibitory gene under sd condition, in other words, the characters of small grain and dwarf of XXA were linked and controlled by a pair of plant height recessive quality gene-rRH.
3. Gene localization of small grain and dwarf characters of XXA:In order to locate rRH gene loci, partial SSR markers of 12 rice chromosomes were selected by PCR that based on BSA, and the semi-dwarf gene of XXA was located to the fifth chromosome. Through intercomparison and calculation, the semi-dwarf gene of XXA located at 8.4 cM upstream from- RM249 in the fifth chromosome, which was different from the sdg downriver from the RM249.
4. breeding male sterile line:Using cross breeding method to transport character of photoperiod sensitive genic male sterile into XXA, then through multi-generation pressurized selecting, a passel of photo-thermo-sensitive genic male sterile lines were gotten; and using methods that backcross, hybrid with wild rice and ionizing radiation, etc. to improve shapes of plant and leaf of XXA and increase its stigma exsertion rate. A passel of male sterile lines with great comprehensive characters were obtained, which were 9sy220,9sy507 and 9d82, etc..
5. Quality combinations with characters of high-yield could be selected:Taking 171S as research object match group widely to determine its combining ability, the results showed that 171S has great combining ability, and quality combinations with characters of high-yield and appropriate growth period could be selected.
6. Preliminary Studies on mixed direct-sowing seed production:Taking 171S as research object and taking restorer line 440, which has similar growth period with 171S's, to conduct mixed direct-sowing seed production; direct sowing experiments were conducted with parents'grain proportion of 1:10,1:20 and 1:30 and repeated three times. The results showed that the average outcrossing rate of 171S was 13.71%, and the differences of outcrossing rates among three levels were not significant. Several factors for low outcrossing rate were analyzed as follows:first, the sensitive degrees of GA3 on parents are different, and the applying amount of GA3 is not easy to control; second, the flag leaves of 171S are long and wide, which affecting pollination; third, the florescence of 171S is not concentrated.
7. Separating machine on the design:Taking XXA as small grain sample to conduct separating experiment for grains and seeds. The results showed that using 3.5 mm circular screen to sift over three times, the small grain and normal large grain (1000-grain weight is around 25 g) could be separated, and the purity of separated seeds reached over 99% that accord with national standards of rice commercial seeds. Based on the separating theory of indented mold-hole, the separating machine had been made by utilizing the different lengths between seeds to ensure the characteristics of mold hole and perfecting related designs, and it has great separating effect.
引文
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