苎麻雄性不育的遗传机理及应用研究
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摘要
苎麻(Boehmeria mvea),又称“中国草”,是我国特有的纤维作物。苎麻雄性不育从上世纪60年代发现至今,已引起了苎麻育种工作者和相关研究者的高度重视。由于对苎麻雄性不育的遗传机制尚不清楚,限制了在育种上利用。因此,确有必要进行深入系统的研究,从而促进苎麻杂交优势的研究和应用。本文采用经典数量遗传、细胞遗传研究方法和现代分子生物学技术手段,通过对苎麻雄性不育的遗传特点、生理生化基础及细胞学特征、雄性不育相关基因的克隆、雄性不育分子标记的建立和杂种优势的利用等方面的研究,对苎麻雄性不育的遗传机理有了较为全面的认识,为利用苎麻雄性不育进行杂交育种奠定了理论基础,指明了合理的应用途径。获得的主要研究结果如下:
1.苎麻雄性不育表型鉴定
通过对苎麻雄性不育及恢复系表型的比较观察,结果表明,相对于正常可育植株,雄性不育主要表现为雌花发育正常,而雄蕾瘦小,不开裂,花药干瘪,内容物很少,无花粉散出;I_2-KI染色无颜色变化,雌花开放时雄蕾逐渐枯萎脱落。
2.苎麻雄性不育的遗传分析
对雄性不育系C26为母本、恢复系B8为父本组配的六个杂交组合的后代植株不育性状分离情况进行调查和分析,表明苎麻雄性不育系C26的育性受一对隐性核基因和细胞质基因共同控制,属于细胞质—核基因互作不育类型,基因型为T(rr)。恢复系B8的育性受一对杂合基因控制,且细胞质与C26不同,基因型为F(Rr)。
3.苎麻雄性不育物质代谢
通过对苎麻雄性不育系和恢复系代谢产物分析后发现,可溶性糖含量在相同的发育阶段,不育系均高于其恢复系,这种差异在大蕾时期最大,不育系C26、C4分别是其恢复系的1.91和2.59倍。淀粉含量随着花蕾发育,不育系无明显变化,而恢复系则不断积累,中蕾时期恢复系B16含量比不育系C4高72.8%,大蕾时期恢复系B8比不育系C26高79.6%。可溶性蛋白质含量随着花蕾发育,不育系和恢复系均逐渐下降,但在材料间表现不一致,不育系C26低于其恢复系B8,而不育系C4则高于其恢复系B16。游离脯氨酸含量的变化,在不育系与恢复系F_1花蕾中可育株含量远高于不育株。
4.苎麻雄性不育同工酶谱分析
通过对苎麻雄性不育系过氧化物酶(POD)和超氧化物歧化酶(SOD)同工酶分析,POD电泳分析未发现雄性不育的特征谱带,但同工酶谱恢复系强于不育系,尤其是迁移率为0.59的条带在各时期均表现如此。SOD电泳分析仅在功能叶中检测到雄性不育系与恢复系在带的强弱上的差异,同样表现恢复系强于不育系,但在雄蕾发育的各时期均无酶带显示。在功能叶、幼蕾、中蕾和大蕾时期,SOD和POD活性均表现恢复系均强于不育系。
5.苎麻雄性不育发生时期分析
通过对苎麻雄性不育系和恢复系的石蜡切片分析,结果显示,雄性不育发生在从造孢细胞到四分体形成的各个阶段,不同的雄性不育系发生不育的时期和败育的原因有所不同。不育系C4主要败育原因为无孢原细胞分化,在发育早期即彻底败育。不育系C26主要败育时期为减数分裂期,败育形式主要为绒毡层巨大化,向内挤压小孢子母细胞,使其不能正常生长发育进入减数分裂,或到后期绒毡层逐渐解体,小孢子母细胞没有进入减数分裂,最终解体。此外还观察到维管束急剧退化现象。
6.苎麻NAD7基因片段的克隆与分析
通过NAD7保守序列设计的引物,应用RT-PCR方法从苎麻雄性不育系C4中克隆了NAD7的cDNA片段,其长度为969 bp,编码322个氨基酸。序列比对表明该cDNA序列及推测的氨基酸序列与拟南芥、油菜有98%的同源性。聚类分析显示,苎麻NAD7基因的进化在植物中处于很古老的地位。
7.苎麻品种(系)与亲本间的遗传关系
从100条ISSR引物中筛选出21条引物,对8份四川苎麻品种(系)及其亲本的遗传关系进行了分析,结果表明,21条引物在8份材料中共扩增出86条带,平均每条引物扩增4.1条带,其中多态性位点71个,各引物扩增出的位点数3～8个,平均可以检测到3.4个多态性位点。聚类分析结果表明,供试品种与其母本遗传距离较远表现特有的偏父本遗传现象。
8.苎麻雄性不育特有分子标记的建立
用ISSR引物U835筛选到1个雄性不育特有的分子标记,对该标记进行克隆测序表明其长度为658bp,并将此标记转化成了稳定的SCAR标记。利用此标记,对强优势组合川苎8号自交群体进行验证,结果显示该标记与雄性不育基因的重组率为3.3%,可用于苎麻雄性不育分子标记辅助育种。
9.苎麻雄性不育系杂种优势利用研究
利用雄性不育材料,进行了杂交组合的测配,对杂交组合的杂种优势、分离变异进行了分析,结果表明苎麻的杂种优势十分显著,强优势主要表现在株高、茎粗及有效株率等方面,为苎麻雄性不育杂种优势的利用提供了科学依据,丰富了苎麻育种理论。
Ramie (Boehmeria. nivea), is an important fiber crops. As our national treasure, it is also called "China grass". Breeding scientists pay a lot of attention to Ramie male sterility since it has been found. However, there are not only few documents and little applications, but also little known of genetic mechanism on Ramie male sterility. To accelerate Ramie breeding process, it is very necessary for us to study Ramie male sterility roundly. In this dissertation, many studies on male-sterility in Ramie were applied, such as genetic analysis, the characteristics in physiology and biochemistry and cytology investigation, NAD7 gene clone and analysis, molecular markers construction and heterosis utilizing. The main results showed as follows:
1. Ramie male sterility phenotype identification
Compared with normally plants, the male sterility plants mainly display on the male flower bud: the male flower bud is thin, small, un-crack, the anther is withered, the contents are very few and without pollen; non-color change with I_2-KI dyes. The female flower opens when the male flower bud withered and gradually falls off. The female flower growth is normal.
2. Ramie male sterility genetic analysis
Genetic analysis of generations of 6 crossed combinations showed that the male sterility character of the male sterility line C26 were controlled by one pair of recessive nuclear genes and cytoplasm gene. So C26, gene type T (rr) ,belong to CMS Fertility. While the gene type of the restoring line B8 was F (Rr).
3. Ramie male sterility material metabolism
In the same developmental stage, the male-sterile line soluble sugar content is higher than its fertility restoring line. The distinct difference was observed in the big flower bud stage: male-sterile line C26 and C4 respective is its fertility restoring line 1.91 and 2.59 times. Along with the flower bud growth, the male-sterile line male flower bud starch content don't changes obviously, but the fertility restoring line accumulates starch steadily: in flower bud stage, fertility restoring line B16 starch content high 72.8% compared to male-sterile line C4, the big flower bud stage, fertility restoring line B8 achieve 79.6% compared to male-sterile line C26. Along with flower bud growth, regardless of the male-sterile line or the fertility restoring line, the soluble protein descends gradually. But with respect to the contents, it displays not consistently between materials: male-sterile line C26 is lower than its restoring line B8, but male-sterile line C4 is higher than its restoring line B16. The free proline content is consistent in the starch content change. In the male-sterile plants and common plants of F1, all material displays the same as its parents, except in the protein content: fertile plant is higher than sterile.
4. Ramie male sterility isozyme analysis
The male sterility characteristic bands have not been observed in the POD electrophoresis. The only difference is the band intensity: the restoring line stronger than the male-sterile line. Therein to Rf 0.59 band in restoring line is stronger than that of male-sterile line in all stages. With respect to SOD, the isozyme bands have not been examined in various times of the male flower bud growth but only in the functional leaves. The mainly difference between male-sterile line and restoring line exist in the band intensity, the same as POD. The POD, SOD activities mensurating results showed that the restoring line stronger than the male-sterile line (fertile plants stronger than sterile plants in F1).
5. Ramie male sterility paraffin section
The paraffin section results indicated that Ramie male sterility occurs in each stage from the spore cell forms to the tetrad, in which male-sterile line C4 main abortion thoroughly in growth early time because of having no spore original cell differentiation, while the male-sterile line C26 main abortion time in meiosis process. The male-sterile line C26 abortion form mainly has: the microspore mother cell cannot develop into meiosis normally, because of the extrusion from the huge tapetum. When the tapetum disintegrates gradually later, the microspore mother cell has still not entered meiosis until disintegrated. Moreover, the phenomenon that canaliculus bundle degenerate suddenly has been found in the experiment.
6. Cloning and analysis of NADH dehydrogenase subunit 7 gene fragment
A cDNA fragment of Ramie NAD7 gene was obtained by RT-PCR with special primers, which were designed according to the conserved sequences of NAD7 gene. The cDNA fragment had the sequence length of 969 bp, encoding 322 amino acids. Analysis of sequence alignment and phylogenetic tree revealed that NAD7 gene of Ramie shared in 98% identity with those of Brassica napus and Arabidopsis thaliana. The results also showed that the NAD7 gene of Ramie is an ancestral type in plant.
7. Genetic relation analysis of Ramie cultivars and their parents from Sichuan with ISSR
21 ISSR primers, which were screened from 100 ISSR primers, were used to analyze and identify the relationship of 8 Ramie with ISSR technology. PCR amplification showed that 86 clear and repeatable bands were obtained in all materials, of which 71 bands appeared to be polymorphic (82.5%). Genetic distance and the cluster analysis indicated that these materials were divided into 2 groups, and the value of genetic distance was ranged from 0.48 to 0.90. Moreover, it was also found that the genetic relationships between hybrids and their parents were not equal, the male parent transfer more genetic loci to their descendants, which indicated that hybrids inherited more genetic information from the male parent.
8. Study on the Molecular Markers Linked to Male Sterile Gene
A specific DNA segment was amplified only in the fertile cultivars but not in the sterile cultivars using the ISSR primer U835. And the ISSR-marker was converted into SCAR marker that can be used to molecular marker-assisted selection of Ramie breeding.
9. Heterosis utilizing of Ramie male sterility
Using the male-sterile breeding materials, the combination of hybrid Ramie male sterility, the combination of hybrid heterosis and separation variation were carried out. The results indicated that heterosis of Ramie male sterile line is quite obvious. Ramie heterosis for the use of CMS not only provides a scientific basi, a new channel of Ramie hybrid and Ramie breeding theory. But also reduce costs, increase farmers income, solve the problems of production, increase Ramie production market competitiveness and resilience of its social and economic efficiency has been remarkable.
1.苎麻雄性不育表型鉴定
通过对苎麻雄性不育及恢复系表型的比较观察,结果表明,相对于正常可育植株,雄性不育主要表现为雌花发育正常,而雄蕾瘦小,不开裂,花药干瘪,内容物很少,无花粉散出;I_2-KI染色无颜色变化,雌花开放时雄蕾逐渐枯萎脱落。
2.苎麻雄性不育的遗传分析
对雄性不育系C26为母本、恢复系B8为父本组配的六个杂交组合的后代植株不育性状分离情况进行调查和分析,表明苎麻雄性不育系C26的育性受一对隐性核基因和细胞质基因共同控制,属于细胞质—核基因互作不育类型,基因型为T(rr)。恢复系B8的育性受一对杂合基因控制,且细胞质与C26不同,基因型为F(Rr)。
3.苎麻雄性不育物质代谢
通过对苎麻雄性不育系和恢复系代谢产物分析后发现,可溶性糖含量在相同的发育阶段,不育系均高于其恢复系,这种差异在大蕾时期最大,不育系C26、C4分别是其恢复系的1.91和2.59倍。淀粉含量随着花蕾发育,不育系无明显变化,而恢复系则不断积累,中蕾时期恢复系B16含量比不育系C4高72.8%,大蕾时期恢复系B8比不育系C26高79.6%。可溶性蛋白质含量随着花蕾发育,不育系和恢复系均逐渐下降,但在材料间表现不一致,不育系C26低于其恢复系B8,而不育系C4则高于其恢复系B16。游离脯氨酸含量的变化,在不育系与恢复系F_1花蕾中可育株含量远高于不育株。
4.苎麻雄性不育同工酶谱分析
通过对苎麻雄性不育系过氧化物酶(POD)和超氧化物歧化酶(SOD)同工酶分析,POD电泳分析未发现雄性不育的特征谱带,但同工酶谱恢复系强于不育系,尤其是迁移率为0.59的条带在各时期均表现如此。SOD电泳分析仅在功能叶中检测到雄性不育系与恢复系在带的强弱上的差异,同样表现恢复系强于不育系,但在雄蕾发育的各时期均无酶带显示。在功能叶、幼蕾、中蕾和大蕾时期,SOD和POD活性均表现恢复系均强于不育系。
5.苎麻雄性不育发生时期分析
通过对苎麻雄性不育系和恢复系的石蜡切片分析,结果显示,雄性不育发生在从造孢细胞到四分体形成的各个阶段,不同的雄性不育系发生不育的时期和败育的原因有所不同。不育系C4主要败育原因为无孢原细胞分化,在发育早期即彻底败育。不育系C26主要败育时期为减数分裂期,败育形式主要为绒毡层巨大化,向内挤压小孢子母细胞,使其不能正常生长发育进入减数分裂,或到后期绒毡层逐渐解体,小孢子母细胞没有进入减数分裂,最终解体。此外还观察到维管束急剧退化现象。
6.苎麻NAD7基因片段的克隆与分析
通过NAD7保守序列设计的引物,应用RT-PCR方法从苎麻雄性不育系C4中克隆了NAD7的cDNA片段,其长度为969 bp,编码322个氨基酸。序列比对表明该cDNA序列及推测的氨基酸序列与拟南芥、油菜有98%的同源性。聚类分析显示,苎麻NAD7基因的进化在植物中处于很古老的地位。
7.苎麻品种(系)与亲本间的遗传关系
从100条ISSR引物中筛选出21条引物,对8份四川苎麻品种(系)及其亲本的遗传关系进行了分析,结果表明,21条引物在8份材料中共扩增出86条带,平均每条引物扩增4.1条带,其中多态性位点71个,各引物扩增出的位点数3～8个,平均可以检测到3.4个多态性位点。聚类分析结果表明,供试品种与其母本遗传距离较远表现特有的偏父本遗传现象。
8.苎麻雄性不育特有分子标记的建立
用ISSR引物U835筛选到1个雄性不育特有的分子标记,对该标记进行克隆测序表明其长度为658bp,并将此标记转化成了稳定的SCAR标记。利用此标记,对强优势组合川苎8号自交群体进行验证,结果显示该标记与雄性不育基因的重组率为3.3%,可用于苎麻雄性不育分子标记辅助育种。
9.苎麻雄性不育系杂种优势利用研究
利用雄性不育材料,进行了杂交组合的测配,对杂交组合的杂种优势、分离变异进行了分析,结果表明苎麻的杂种优势十分显著,强优势主要表现在株高、茎粗及有效株率等方面,为苎麻雄性不育杂种优势的利用提供了科学依据,丰富了苎麻育种理论。
Ramie (Boehmeria. nivea), is an important fiber crops. As our national treasure, it is also called "China grass". Breeding scientists pay a lot of attention to Ramie male sterility since it has been found. However, there are not only few documents and little applications, but also little known of genetic mechanism on Ramie male sterility. To accelerate Ramie breeding process, it is very necessary for us to study Ramie male sterility roundly. In this dissertation, many studies on male-sterility in Ramie were applied, such as genetic analysis, the characteristics in physiology and biochemistry and cytology investigation, NAD7 gene clone and analysis, molecular markers construction and heterosis utilizing. The main results showed as follows:
1. Ramie male sterility phenotype identification
Compared with normally plants, the male sterility plants mainly display on the male flower bud: the male flower bud is thin, small, un-crack, the anther is withered, the contents are very few and without pollen; non-color change with I_2-KI dyes. The female flower opens when the male flower bud withered and gradually falls off. The female flower growth is normal.
2. Ramie male sterility genetic analysis
Genetic analysis of generations of 6 crossed combinations showed that the male sterility character of the male sterility line C26 were controlled by one pair of recessive nuclear genes and cytoplasm gene. So C26, gene type T (rr) ,belong to CMS Fertility. While the gene type of the restoring line B8 was F (Rr).
3. Ramie male sterility material metabolism
In the same developmental stage, the male-sterile line soluble sugar content is higher than its fertility restoring line. The distinct difference was observed in the big flower bud stage: male-sterile line C26 and C4 respective is its fertility restoring line 1.91 and 2.59 times. Along with the flower bud growth, the male-sterile line male flower bud starch content don't changes obviously, but the fertility restoring line accumulates starch steadily: in flower bud stage, fertility restoring line B16 starch content high 72.8% compared to male-sterile line C4, the big flower bud stage, fertility restoring line B8 achieve 79.6% compared to male-sterile line C26. Along with flower bud growth, regardless of the male-sterile line or the fertility restoring line, the soluble protein descends gradually. But with respect to the contents, it displays not consistently between materials: male-sterile line C26 is lower than its restoring line B8, but male-sterile line C4 is higher than its restoring line B16. The free proline content is consistent in the starch content change. In the male-sterile plants and common plants of F1, all material displays the same as its parents, except in the protein content: fertile plant is higher than sterile.
4. Ramie male sterility isozyme analysis
The male sterility characteristic bands have not been observed in the POD electrophoresis. The only difference is the band intensity: the restoring line stronger than the male-sterile line. Therein to Rf 0.59 band in restoring line is stronger than that of male-sterile line in all stages. With respect to SOD, the isozyme bands have not been examined in various times of the male flower bud growth but only in the functional leaves. The mainly difference between male-sterile line and restoring line exist in the band intensity, the same as POD. The POD, SOD activities mensurating results showed that the restoring line stronger than the male-sterile line (fertile plants stronger than sterile plants in F1).
5. Ramie male sterility paraffin section
The paraffin section results indicated that Ramie male sterility occurs in each stage from the spore cell forms to the tetrad, in which male-sterile line C4 main abortion thoroughly in growth early time because of having no spore original cell differentiation, while the male-sterile line C26 main abortion time in meiosis process. The male-sterile line C26 abortion form mainly has: the microspore mother cell cannot develop into meiosis normally, because of the extrusion from the huge tapetum. When the tapetum disintegrates gradually later, the microspore mother cell has still not entered meiosis until disintegrated. Moreover, the phenomenon that canaliculus bundle degenerate suddenly has been found in the experiment.
6. Cloning and analysis of NADH dehydrogenase subunit 7 gene fragment
A cDNA fragment of Ramie NAD7 gene was obtained by RT-PCR with special primers, which were designed according to the conserved sequences of NAD7 gene. The cDNA fragment had the sequence length of 969 bp, encoding 322 amino acids. Analysis of sequence alignment and phylogenetic tree revealed that NAD7 gene of Ramie shared in 98% identity with those of Brassica napus and Arabidopsis thaliana. The results also showed that the NAD7 gene of Ramie is an ancestral type in plant.
7. Genetic relation analysis of Ramie cultivars and their parents from Sichuan with ISSR
21 ISSR primers, which were screened from 100 ISSR primers, were used to analyze and identify the relationship of 8 Ramie with ISSR technology. PCR amplification showed that 86 clear and repeatable bands were obtained in all materials, of which 71 bands appeared to be polymorphic (82.5%). Genetic distance and the cluster analysis indicated that these materials were divided into 2 groups, and the value of genetic distance was ranged from 0.48 to 0.90. Moreover, it was also found that the genetic relationships between hybrids and their parents were not equal, the male parent transfer more genetic loci to their descendants, which indicated that hybrids inherited more genetic information from the male parent.
8. Study on the Molecular Markers Linked to Male Sterile Gene
A specific DNA segment was amplified only in the fertile cultivars but not in the sterile cultivars using the ISSR primer U835. And the ISSR-marker was converted into SCAR marker that can be used to molecular marker-assisted selection of Ramie breeding.
9. Heterosis utilizing of Ramie male sterility
Using the male-sterile breeding materials, the combination of hybrid Ramie male sterility, the combination of hybrid heterosis and separation variation were carried out. The results indicated that heterosis of Ramie male sterile line is quite obvious. Ramie heterosis for the use of CMS not only provides a scientific basi, a new channel of Ramie hybrid and Ramie breeding theory. But also reduce costs, increase farmers income, solve the problems of production, increase Ramie production market competitiveness and resilience of its social and economic efficiency has been remarkable.
引文
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