水稻育性调控的分子遗传研究进展

杂交水稻为世界粮食安全做出了重要贡献。细胞质雄性不育和光/温敏不育分别是三系和两系杂交稻生产利用的遗传基础,而(亚)种间杂种不育则是杂交稻生产中要克服的主要技术瓶颈。因此,水稻育性调控是杂交水稻生产技术的关键,也是研究植物核质互作和物种生殖隔离等基础科学问题的重要模型。我国植物遗传学家在阐明杂交水稻育性调控的分子遗传基础领域做出了重要贡献。本文回顾了我国杂交水稻的发展历程,系统总结了杂交水稻生产涉及的细胞质雄性不育与恢复、光/温敏不育与育性转换、杂种不育与亲和性的遗传基础和分子作用机制,探讨了我国杂交水稻生产存在的问题,指明了水稻杂种优势利用的发展方向。     

从水稻同源三倍体中鉴定出无融合生殖种质TAR

水稻杂种优势的成功应用使水稻单产提高了20%,这主要得益于70年代初细胞质雄性不育系的发现和利用。但传统的三系法杂交水稻必须做到不育系、保持系、恢复系“三系”配套,选育程序和杂交种子生产环节较复杂,以致培育目标组合的周期长、效率低、推广环节多,同时种子成本高、价格贵[1]。80年代光敏核不育水稻以其“一系两用、配组自由”等优越性大有取代三系杂交水稻的趋势。但是,光敏核不育特性同时又受温度的调控,加之育性波动等缺陷使两系杂交水稻的研究和利用受到严重制约[2]。80年代后期,科技工作者开始探索利用无融合生殖固定水稻杂种优…     

杂交水稻及其三系对日照长度的反应

杂交水稻的培育成功,是农业生产上的一项突破,使粮食大幅度增产。水稻是喜温的短日照植物,光照阶段的发育特性是确定水稻适应地区、安排播种季节、制订田间管理技术措施的依据之一。要使杂交水稻充分发挥其优势,在栽培上必须要按照杂交水稻的生长发育规律进行。我们于1977至1979年的三年中,对目前推广的及准备推广的杂交水稻和它们的不育系及恢复系对日照长度的反应进行了试验研究。     

利用生物技术创建主要作物雄性不育杂交育种和制种的技术体系

雄性不育技术在作物杂种优势利用和杂交种生产中发挥着重要作用。基于核质互作雄性不育的“三系法”与光温敏核不育的“两系法”已经在水稻等主要作物的杂交制种中获得了广泛应用,但是存在着资源利用效率低、育性不稳定、易受外界环境影响等诸多问题。近三十年来,利用生物技术创建不同类型的植物雄性不育系取得了一系列突破性进展。主要针对玉米、水稻、小麦三大作物的基因工程雄性不育技术的最新进展进行总结,特别详细地描述了本实验室最近研究创制的玉米多控不育技术体系,以期为相关研究和产业化应用提供技术参考。     

两系法杂交水稻

七十年代初我国学者发现了“野败”细胞质和IR系统的恢复基因,成功地培育出了核质互作籼型杂交水稻,使得水稻产量有了大幅度提高。核质互作型杂交水稻是基于不育系、保持系、恢复系的配套实现不育系的繁殖、杂交种子(F_1)的生产的,因此三系法育种程序和生产环节复杂,选育新组合的周期长、效率低,同时杂种种子有成本高、产量低,这些都是三系杂交稻在种植面积上受到限制的原因。     

印水型杂交水稻开创杂交水稻超高产制种新时代

在2005年国家主推的19个籼型超级杂交稻中有9个印水型杂交水稻,这种水稻连续创造了世界水稻最高产量纪录, 1／15hm2产达到了1 231．17 kg。印水型杂交水稻由中国水稻研究所主持完成,其育成和推广,从制种产量、杂种产量和米质等方面把我国杂交水稻的生产水平总体提高到一个新台阶,开创了杂交水稻超高产制种新时代,米质主要指标达到国际3-2级;制种产量创造并保持了我国和世界制种最高产量纪录,达到每1／15hm2440 kg;制种成本降低50％以上。从1998年至2004年,全国累计推广印水型杂交水稻24万hm2以上,增产稻谷64亿kg,创造经济效益 95亿元。该所“印水型水稻不育胞质的发掘及应用”项目荣获了2005年度国家科技进步一等奖和浙江省科技进步一等奖。     

水稻CMS不育系细胞质遗传效应的研究进展

在三系杂交水稻生产中,不育系的细胞质对杂种一代的表现具有一定的效应。同一种不育细胞质对不同性状的效应是不同的;不同的不育细胞质对同一性状的效应也存在着差异;且对大多数性状表现为负向效应。可通过扩大不育细胞质源以选用优质细胞质或选用强优恢复系来减少或消除其负效应,培育同核异质的多胞质不育系来适应杂交水稻生产的需要。     

水稻CMS不育系细胞质遗传效应的研究进展

在三系杂交水稻生产中,不育系的细胞质对杂种一代的表现具有一定的效应。同一种不育细胞质对不同性状的效应是不同的;不同的不育细胞质对同一性状的效应也存在着差异;且对大多数性状表现为负向效应。可通过扩大不育细胞质源以选用优质细胞质或选用强优恢复系来减少或消除其负效应,培育同核异质的多胞质不育系来适应杂交水稻生产的需要。     

环境对杂交水稻胞质及核质互作遗传表现的影响─Ⅳ．不同N条件下不育胞质对杂种1代多胺代谢的影响

研究结果表明,杂交水稻在不适宜的Ｎ条件下,ＡＤＣ活性及其代谢产物Ｐｕｔ含量和Ｓｐｍ／Ｓｐｄ值均有显著提高,其幅度随核质组成及其与环境互作的不同而异．不育胞质对杂交水稻ＡＤＣ活性的影响呈稳定的正效应,环境并不改变不育胞质对其作用的性质,但可调节其作用强度．在适宜的生理阈值内,通过栽培措施提高上述各生理指标有利于促进不育胞质对杂交水稻产量性状的正向影响和提高杂种优势强度．而在不适宜条件下上述各生理指标则超越其生理阈值．不利于水稻本身的生长发育．此外,杂交水稻在不适宜Ｎ条件下大量积累Ｐｕｔ,往往伴随着ＩＡＡｏｓｅ和ＤＡＯ活性的提高和ＩＡＡ含量下降．     

新版教材中问题解答3则

1　什么叫两系法杂交水稻我国是世界上第 1个将杂交优势应用于水稻生产的国家。袁隆平于 196 4年开始水稻杂交优势的应用研究 ,于 1973年实现籼型三系 (不育系、保持系和恢复系 )配套 ,而两系法杂交水稻中的不育系则是一种光敏型核雄性不育类型。所谓光敏型核雄性不育是指雄蕊育性对光照敏感 ,在夏季长日照下表现为雄性不育 ,在秋季短日照下雄性又可育。有研究表明 ,在短日照下光感核不育基因不表达 ,但在穗分化期的任何阶段给予长日照 ,该基因即可表达 ,只有从第 2次枝梗原始体分化期开始 ,持续给予长日照花粉才彻底败育 ,由此可见从第 2次…     

Targeted mutagenesis of a conserved anther‐expressed P450 gene confers male sterility in monocots

Targeted mutagenesis using programmable DNA endonucleases has broad applications for studying gene function in planta and developing approaches to improve crop yields. Recently, a genetic method that eliminates the need to emasculate the female inbred during hybrid seed production, referred to as Seed Production Technology, has been described. The foundation of this genetic system relied on classical methods to identify genes critical to anther and pollen development. One of these genes is a P450 gene which is expressed in the tapetum of anthers. Homozygous recessive mutants in this gene render maize and rice plants male sterile. While this P450 in maize corresponds to the male fertility gene Ms26, male fertility mutants have not been isolated in other monocots such as sorghum and wheat. In this report, a custom designed homing endonuclease, Ems26+, was used to generate in planta mutations in the rice, sorghum and wheat orthologs of maize Ms26. Similar to maize, homozygous mutations in this P450 gene in rice and sorghum prevent pollen formation resulting in male sterile plants and fertility was restored in sorghum using a transformed copy of maize Ms26. In contrast, allohexaploid wheat plants that carry similar homozygous nuclear mutations in only one, but not all three, of their single genomes were male fertile. Targeted mutagenesis and subsequent characterization of male fertility genes in sorghum and wheat is an important step for capturing heterosis and improving crop yields through hybrid seed.     

Male sterility systems in wheat and opportunities for hybrid wheat development

The common wheat (Triticum aestivum L.) is a poly(hexa)ploid, derived from an amphi-diploidization process involving the donor species—Triticum urartu, Aegilops speltoides, Triticum turgidum, and Aegilops tauschii. The genetic diversity of the autogamous wheat is narrow, which is a major reason for lesser rate of yield gain in wheat, in contrast to rice and maize. It is desirable to encourage hybrid breeding, i.e., combining different lines into genetically divergent heterotic pools. Thus, hybrid plants are a unique combination of desired alleles produced by crossing between genetically different parental lines. Hybrid seed production in a crop requires male-sterile female parents along with a reliable outcrossing system. The male-sterile female parent prevents pollen shedding and self-fertilization, maintaining the purity of hybrid seeds. An outcrossing system enhances hybrid seed production. This article emphasizes the biological relevance of crossbreeding and self-pollination in wheat, and reviews different male sterility systems which could be utilized for the development of hybrid wheat. Several biotechnological approaches and their practical utility in generating cross-compatible male-sterile female parent lines have been discussed.     

No effect of transgene and strong wild parent effects on seed dormancy in crop–wild hybrids of rice: implications for transgene persistence in wild populations

Soil seed banks act as a gene pool for local plant species and, as such, can buffer local populations, especially those experiencing challenging environmental conditions. Seed dormancy has important implications to dynamics of soil seed banks. Therefore, estimating the seed dormancy of transgenic crop–wild hybrids could shed light on the persistence of transgenes in wild‐plant soil seed banks. Individuals from eight populations of wild rice Oryza rufipogon were crossed with those of three insect‐resistant transgenic rice lines. Selfed (F2–F4) and backcrossed populations (BC1, BC1F2 and BC1F3) were then made from the hybrids. Seed germination was tested under three treatments: (a) normal; (b) overwintering in soil; and (c) one‐week heat‐shocking. The effects of transgene, wild parent and hybrid generation on hybrid seed germination were examined. No significant effect of insect‐resistant transgenes (Bt and CpTI) was detected on the seed dormancy of crop–wild hybrids, while a significant wild parent effect was found. The seeds of advanced generation hybrids have higher germination percentages and lower dormancy than do those of F1 and BC1 generations. The study showed that the dormancy of hybrid seeds was determined mainly by their genetic backgrounds. All hybrid seeds have higher germination percentages and lower dormancy (and, consequently, a poorer overwintering ability), compared with wild seeds, and reduce dormancy would contribute to a fitness disadvantage, compared with wild types. Therefore, such seeds might form part of naturally occurring soil seed banks, through which crop genes would persist in wild populations.     

The Genetic and Biochemical Mechanisms Underlying Cereal Seed Dormancy

The crop seeds have been a staple food for humans, and seed yield is important for sustaining agriculture development and enhancing human adaptability to food risks. The phenomenon of pre-harvest sprouting (PHS), caused by seed dormancy deficiency, and the phenomenon of low seedling emergence caused by seed deep dormancy, will lead to a reduction in agricultural production. Therefore, it is particularly important to understand the regulation mechanisms of seed dormancy. There are many studies on the regulation of seed dormancy in rice, but there are few studies on the regulation of seed dormancy in other crops, and the research on its mechanism is not thorough enough. In this paper, we comprehensively summarize the regulation mechanisms of cereal seed dormancy, including rice, barley and wheat, discussing the integral mechanism of seed dormancy. This information should provide new insights for developing versatile cultivated lines to improve crop yield and economic benefits.     

A straight-forward seed production technology system for foxtail millet (Setaria italica)

For autogamous crops, a precondition for using heterosis is to produce sufficient pure male-sterile female parents that can be used to produce hybrid seeds. To date, cytoplasmic male sterility(CMS)and environment-sensitive genic male sterility(EGMS) have been used commercially to exploit heterosis for autogamous species. However, neither CMS nor EGMS has been established for foxtail millet(Setaria italica). Here, we report on the establishment and application of a seed production technology(SPT)...     

Improving panicle exsertion of rice cytoplasmic male sterile line by combination of artificial microRNA and artificial target mimic

The adoption of hybrid rice caused the second leap in rice yield after the ‘green revolution’ and contributes substantially to food security of China and the world. However, almost all cytoplasmic male sterile lines (A lines) as females of hybrid rice have a natural deficiency of ‘panicle enclosure’, which blocks pollination between the A line and the fertility restorer line as the male (R line) of hybrid rice and decreases seed yield. In hybrid rice seed production, exogenous ‘920’ (the active ingredient is gibberellin A₃) must be applied to eliminate or alleviate panicle enclosure of the A line; however, this not only increases production cost and pollutes the environment, it also decreases seed quality. In this study, we designed a transgenic approach to improve plant height and panicle exsertion of the A line to facilitate hybrid rice production and maintain the semi‐dwarf plant type of the hybrid. This approach comprising two components—artificial microRNA (amiRNA) and artificial target mimicry—can manipulate the differential expression of the endogenous Eui1 gene that is associated with rice internode elongation in the A line and the hybrid. amiRNA is a recently developed gene silencing method with high specificity, while target mimicry is a natural mechanism inhibiting the miRNA function that was also recently characterized. This approach provides a paradigm to tune the expression of endogenous genes to achieve the desired phenotype by combining amiRNA and artificial target mimicry technologies.     

Strategy and utilization of a herbicide resistance gene in two-line hybrid rice

Hybrid rice plays an important role in China's aim to improve rice production as it accounts for some 50% of rice planting area but produces about 60% of the total rice grain. However, the existing three-line system used in hybrid rice production has its limitations. The two-line system, which makes use of photoperiod-sensitive genic male-sterile (PGMS) and thermo-sensitive genic male-sterile (TGMS) lines to generate the male-sterile parental line, was developed to overcome some of these limitations. The sterility of the male-sterile line of two-line hybrid rice, however, fluctuates when the temperature-sensitive phase of fertility encounters abnormal low temperatures during hybrid seed production, which induces selfing and decreases the purity of hybrid. We describe here the strategy of utilizing a herbicide resistance gene in two-line hybrid rice to eliminate this fluctuation in the sterility of the P/TGMS lines during hybrid seed production and reports the development of the herbicide resistance restorer line Bar68-1 and its herbicide-resistant early season hybrid rice Xiang125s/Bar68-1. When the restorer line and its derived hybrid are herbicide resistant, the selfed seeds can be removed easily from the hybrid by herbicide spraying. A herbicide resistance gene bar was transferred into a restorer line by particle bombardment. The resulting transgenic restorer line Bar68-1 and its hybrid Xiang125 s/Bar68-1 inherited stable herbicide resistance. The purity of Xiang125s/Bar68-1 was increased by spraying the seed bed with herbicide, which resulted in a significant increase in yield, grain quality, and disease resistance in comparison to the controls in a regional trial.     

大豆细胞核雄性不育基因研究进展

雄性不育是指植物雄蕊不能正常生长和产生有活力花粉粒的现象。利用雄性不育突变体开展杂交育种工作,是快速提高作物单产的有效途径。目前,通过杂种制种已大幅度提高了水稻(Oryza sativa L.)、玉米(Zea mays L.)和小麦(Triticum aestivum L.)等作物的产量。大豆(Glycinemax(L.)Merr.)作为自花授粉作物,通过人工去雄生产杂交种子不仅困难而且经济上不可行。由于适用于杂交种生产的不育系资源短缺,目前大豆还没有实现大规模杂种优势利用。因此,快速实现大豆杂种优势利用迫切需要鉴定稳定的大豆雄性不育系统。本文总结了大豆细胞核雄性不育(genic male sterility, GMS)突变体及不育基因研究进展,同时结合拟南芥(Arabidopsis thaliana)、水稻和玉米中已报道的细胞核雄性不育基因,从反向遗传学的角度,为大豆核雄性不育基因的鉴定提供依据。     

Hybrid seed production and the challenge of propagating male-sterile plants

The introduction of hybrid crop varieties has enabled spectacular increases in productivity owing to hybrid vigor and increased uniformity. To produce hybrid seeds, a pollination control system is required to prevent unwanted self-pollination. In crop species with hermaphrodite flowers, this can be a major challenge. Over the past decade, new pollination control systems have been developed with the aid of genetic engineering, mainly based on the generation of nuclear-encoded male sterility. The successful application of these systems for large-scale hybrid seed production depends on whether the male-sterile female parent line can be multiplied efficiently and economically. In spite of its relevance, the propagation of the male-sterile line has often been overlooked in the development of pollination control systems.