崛起的生物学科生长点──无融合生殖学

无融合生殖是与有性生殖、无性生殖并存的三大生殖系统之一。自从１７４５年发现弧雄生殖现象以来,关于无融合生殖的分类、形态结构、胚胎发生、遗传进化、生理生化等方面的研究不断开展,已从３６个科３００多种植物中发现无融合生殖现象。特别是八十年代以来,无融合生殖研究已成为生物学科的新热点。全世界几十个国家,２００多个实验室正在从事这一研究。国际性无融合生殖研究协作网已经成立。国际性学术会议相继召开。关于无融合生殖的专著已经出版,专门杂志也已问世。这些事实说明,无融合生殖学这个生物学科的生长点已经崛起。但是由于无融合生殖所具有的特殊性给研究带来许多困难,给生物学科的诸多领域留下了不少空白点和世纪性难题。有关无融合生殖的概念和范围,无融合生殖的遗传,无融合生殖是进化还是退化,无融合生殖在胚胎学、生殖生物学、发育生物学中的重大问题,如：为何无融合生殖植物大小孢子母细胞分裂行为不一致？二倍体孢子生殖中大孢子母细胞胼胝质缺乏是无融合生殖的因还是果？怎样理解同一珠心组织中不同细胞分化？无融合生殖胆和胚乳的形成机制和相互关系以及在作物育种中应用的诱人前景都是当前研究的热门话题。本文都进行了必要的介绍和讨论。     

植物无融合生殖研究新进展

无融合生殖是指不经过雌雄配子融合而产生种子的一种特殊生殖方式,能使基因型的杂合性得以保持,从而可以固定杂种优势,对作物育种具有极其重要的意义。目前大量的研究都在设法将无融合生殖作为一种重要的植物育种手段。本文对近几年来无融合生殖新种质资源的发现、主要研究方法、遗传机制和相关基因等方面的最新进展作了介绍,并对无融合生殖研究中存在的问题和发展前景作了讨论。     

雾灵山草地早熟禾多胚囊和多胚的研究

无融合生殖是指未经炷卵事例而产生后代的特殊生殖方式,它可以分为单倍体无融合生殖和二倍体无融合生殖;对于作物意义更大的是二倍体无融合生殖。多胚囊和多胚现象ＳＨＩ是无融合生殖的表现形式。本文运用石蜡切片法、子房整体透明法研究了雾灵山草地早熟禾（Ｐｏａ ｐｒａｔｅｎｓｉｓ Ｌ．）多胚囊和多胚现象。结果表明,（１）草地早熟禾多胚囊来源有两种：一是自大孢子母细胞,二是来自珠心细胞;（２）草地早熟禾多 来源有     

雾灵山草地早熟禾多胚囊和多胚现象的研究

无融合生殖是指未经精卵融合而产生后代的特殊生殖方式,它可以分为单倍体无融合生殖和二倍体无融合生殖;对于作物改良意义更大的是二倍体无融合生殖。多胚囊和多胚现象SHI是无融合生殖的表现形式。本文运用石蜡切片法、子房整体透明法研究了雾灵山草地早熟禾〖WTBX〗（Poa pratensis〖WTBZ〗 L.）多胚囊和多胚现象。结果表明,（1）草地早熟禾多胚囊来源有两种：一是来自大孢子母细胞,二是来自珠心细胞;（2）草地早熟禾多胚来源有四个：其一是有性生殖胚,其二是孤雌生殖胚,其三是无配子生殖胚,其四是珠心胚。     

单子叶植物无融合生殖的研究进展

植物的无融合生殖是指不经过雌雄配子融合而产生种子的一种特殊生殖方式。由于利用无融合生殖途径可以固定杂种优势,从而改良现有植物的育种策略,因此对无融合生殖的研究已成为生物学科的新生长点。本文主要从无融合生殖的概念和类型,无融合生殖在单子叶植物中的分布,无融合生殖的胚胎学,分子生物学和遗传学机制及创造新的无融合生殖种质资源的方法等6方面对单子叶植物的无融合生殖的研究进展进行了综述,并提出了今后开展无融合生殖研究的思路和设想。     

单子叶植物无融合生殖的研究进展

植物的无融合生殖是指不经过雌雄配子融合而产生种子的一种特殊生殖方式。由于利用无融合生殖途径可以固定杂种优势 ,从而改良现有植物的育种策略 ,因此对无融合生殖的研究已成为生物学科的新生长点。本文主要从无融合生殖的概念和类型 ,无融合生殖在单子叶植物中的分布 ,无融合生殖的胚胎学 ,分子生物学和遗传学机制及创造新的无融合生殖种质资源的方法等 6方面对单子叶植物的无融合生殖的研究进展进行了综述 ,并提出了今后开展无融合生殖研究的思路和设想     

植物无融合生殖研究进展

植物无融合生殖是一种特殊的无性生殖方式 ,它不经过精卵融合即可繁殖后代 ,其二倍体子代基因型与母本精确相同 ,可以固定杂种优势 ,对于作物育种等工作具有巨大的经济意义。对无融合生殖的分类、遗传进化、发生机制、分子机理等方面进行了介绍。并对无融合生殖的一些最新的研究进展 :无孢子生殖专化基因组区、脱调节理论、基因组冲撞观点、表观遗传基因调节理论等进行了简要的评述。并简单介绍了无融合生殖甜菜单体附加系目前的研究进展 。     

植物无融合生殖研究进展北大核心CSCD

无融合生殖是一种不发生雌雄配子核融合而产生种子的一种无性繁殖过程。有些无融合生殖产生的种子是其母本的克隆,可以保留母本的基因型,因此无融合生殖可用于杂种优势的固定。尽管无融合生殖具有潜在的应用价值,但其形成机理十分复杂,表现在无融合生殖有多种表现形式,且受控的途径多样,遗传机制复杂,至今尚无定论,研究方法也多种多样。对近年来无融合生殖研究方面取得的进展进行了概述,旨在为深入研究无融合生殖提供参考。     

植物无融合生殖研究进展

无融合生殖是一种不发生雌雄配子核融合而产生种子的一种无性繁殖过程。有些无融合生殖产生的种子是其母本的克隆,可以保留母本的基因型,因此无融合生殖可用于杂种优势的固定。尽管无融合生殖具有潜在的应用价值,但其形成机理十分复杂,表现在无融合生殖有多种表现形式,且受控的途径多样,遗传机制复杂,至今尚无定论,研究方法也多种多样。对近年来无融合生殖研究方面取得的进展进行了概述,旨在为深入研究无融合生殖提供参考。     

禾本科植物无融合生殖(综述)

禾本科植物包含了世界上最重要的农作物,也包含了最多的无融合生殖的种类,通过无融合生殖可将农作物的F1代杂种优势固定下来,这在固定农作物杂种优势的利用上具有巨大的潜力,然而禾本科植物无融合生殖作为其繁殖多样性的一种形式,在系统进化过程中的作用是非常复杂的,本文统计了禾本科无融合生殖的分布,概述了其无融合生殖的细胞学,遗传学和分子生物学研究进展。     

Molecular and genetic regulation of apomixis

Apomixis is defined as the asexual plant reproduction through seeds that results in the production of genetically uniform progeny. In fact, apomixis could be considered as a natural way of cloning. Currently there are more than 400 plant species known to use apomixis as a strategy for their propagation. The primary fundamental aspects of apomixis are the bypassing of meiosis and parthenogenetic development of the embryo without fertilization Apomixis attracts special attention because of its potential value for agriculture, as it could be harnessed for plant breeding programs enabling the permanent fixation of heterosis in crop plants. A better understanding of the molecular and genetic regulation of apomixis is important for developmental and evolutionary perspectives but also for implementation of engineering of apomixis traits into agricultural crop plants. Despite apomixis is considered as one of the key technologies for the improving agriculture, it is currently not fully known how the genetic and molecular regulation of this important trait occurs. In this review, an up to date information on the biology of apomixis and the known genes and genetic loci associated with regulation of different components of apomixis is provided.     

Apomixis technology and the paradox of sex

Most plant species produce genetically variable seeds by the fusion of meiotically reduced egg cells and pollen grains. However, a small proportion of seed plants produces clonal, asexual seeds by the process of apomixis. The fixation of heterosis by apomixis is of great interest for plant breeding. The prospect of changing sexual crop species into apomictic crop species by genetic engineering--apomixis technology--has recently caused a boom in apomixis research. According to evolutionary biological theories, a dominant apomixis gene will rapidly become fixed in an outcrossing sexual population. Therefore, in theory, apomixis transgenes could have unconditional advantages that could result in the uncontrollable spread of the transgenes. By contrast, 'classic' transgenes might only have conditional advantages. Paradoxically, sexual reproduction and not apomixis is common in nature. However, this is no guarantee that apomixis transgenes will be ecologically safe because there could be essential differences between natural and transgenic apomicts.     

Developmental genetics of gametophytic apomixis

Some higher plants reproduce asexually by apomixis, a natural way of cloning through seeds. Apomictic plants produce progeny that are an exact genetic replica of the mother plant. The replication is achieved through changes in the female reproductive pathway such that female gametes develop without meiosis and embryos develop without fertilization. Although apomixis is a complex developmental process, genetic evidence suggests that it might be inherited as a simple mendelian trait - a paradox that could be explained by recent data derived from apomictic species and model sexual organisms. The data suggest that apomixis might rely more on a global deregulation of sexual reproductive development than on truly new functions, and molecular mechanisms for such a global deregulation can be proposed. This new understanding has direct consequences for the engineering of apomixis in sexual crop species, an application that could have an immense impact on agriculture.     

Apomixis technology development-virgin births in farmers' fields?

Apomixis is the process of asexual reproduction through seed, in the absence of meiosis and fertilization, generating clonal progeny of maternal origin. Major benefits to agriculture could result from harnessing apomixis in crop plants. Although >400 apomictic plant species are known, apomixis is rare among crop plants, and the transfer of apomixis to crop varieties by conventional breeding has been largely unsuccessful. Because apomictic and sexual pathways are closely related, de novo engineering of apomixis might be achieved in sexually reproducing crops. Early consideration of issues relating to biosafety and intellectual property (IP) management can facilitate the acceptance and deployment of apomixis technology in agriculture.     

Apomixis for crop improvement

Summary Apomixis is a genetically controlled reproductive process by which embryos and seeds develop in the ovule without female meiosis and egg cell fertilization. Apomixis produces seed progeny that are exact replicas of the mother plant. The major advantage of apomixis over sexual reproduction is the possibility to select individuals with desirable gene combinations and to propagate them as clones. In contrast to clonal propagation through somatic embryogenesis or in vitro shoot multiplication, apomixis avoids the need for costly processes, such as the production of artificial seeds and tissue culture. It simplifies the processes of commercial hybrid and cultivar production and enables a large-scale seed production economically in both seed- and vegetatively propagated crops. In vegetatively reproduced plants (e.g., potato), the main applications of apomixis are the avoidance of phytosanitary threats and the spanning of unfavorable seasons. Because of its potential for crop improvement and global agricultural production, apomixis is now receiving increasing attention from both scientific and industrial sectors. Harnessing apomixis is a major goal in applied plant genetic engineering. In this regard, efforts are focused on genetic and breeding strategies in various plant species, combined with molecular methods to analyze apomictic and sexual modes of reproduction and to identify key regulatory genes and mechanisms underlying these processes. Also, investigations on the components of apomixis, i.e., apomeiosis, parthenogenesis, and endosperm development without fertilization, genetic screens for apomictic mutants and transgenic approaches to modify sexual reproduction by using various regulatory genes are receiving a major effort. These can open new avenues for the transfer of the apomixis trait to important crop species and will have far-reaching potentials in crop improvement regarding agricultural production and the quality of the products.     

Apomixis in plant reproduction: a novel perspective on an old dilemma

Seed is one of the key factors of crop productivity. Therefore, a comprehension of the mechanisms underlying seed formation in cultivated plants is crucial for the quantitative and qualitative progress of agricultural production. In angiosperms, two pathways of reproduction through seed exist: sexual or amphimictic, and asexual or apomictic; the former is largely exploited by seed companies for breeding new varieties, whereas the latter is receiving continuously increasing attention from both scientific and industrial sectors in basic research projects. If apomixis is engineered into sexual crops in a controlled manner, its impact on agriculture will be broad and profound. In fact, apomixis will allow clonal seed production and thus enable efficient and consistent yields of high-quality seeds, fruits, and vegetables at lower costs. The development of apomixis technology is expected to have a revolutionary impact on agricultural and food production by reducing cost and breeding time, and avoiding the complications that are typical of sexual reproduction (e.g., incompatibility barriers) and vegetative propagation (e.g., viral transfer). However, the development of apomixis technology in agriculture requires a deeper knowledge of the mechanisms that regulate reproductive development in plants. This knowledge is a necessary prerequisite to understanding the genetic control of the apomictic process and its deviations from the sexual process. Our molecular understanding of apomixis will be greatly advanced when genes that are specifically or differentially expressed during embryo and embryo sac formation are discovered. In our review, we report the main findings on this subject by examining two approaches: i) analysis of the apomictic process in natural apomictic species to search for genes controlling apomixis and ii) analysis of gene mutations resembling apomixis or its components in species that normally reproduce sexually. In fact, our opinion is that a novel perspective on this old dilemma pertaining to the molecular control of apomixis can emerge from a cross-check among candidate genes in natural apomicts and a high-throughput analysis of sexual mutants.     

High-resolution physical mapping in Pennisetum squamulatum reveals extensive chromosomal heteromorphism of the genomic region associated with apomixis

Gametophytic apomixis is asexual reproduction as a consequence of parthenogenetic development of a chromosomally unreduced egg. The trait leads to the production of embryos with a maternal genotype, i.e. progeny are clones of the maternal plant. The application of the trait in agriculture could be a tremendous tool for crop improvement through conventional and nonconventional breeding methods. Unfortunately, there are no major crops that reproduce by apomixis, and interspecific hybridization with wild relatives has not yet resulted in commercially viable germplasm. Pennisetum squamulatum is an aposporous apomict from which the gene(s) for apomixis has been transferred to sexual pearl millet by backcrossing. Twelve molecular markers that are linked with apomixis coexist in a tight linkage block called the apospory-specific genomic region (ASGR), and several of these markers have been shown to be hemizygous in the polyploid genome of P. squamulatum. High resolution genetic mapping of these markers has not been possible because of low recombination in this region of the genome. We now show the physical arrangement of bacterial artificial chromosomes containing apomixis-linked molecular markers by high resolution fluorescence in situ hybridization on pachytene chromosomes. The size of the ASGR, currently defined as the entire hemizygous region that hybridizes with apomixis-linked bacterial artificial chromosomes, was estimated on pachytene and mitotic chromosomes to be approximately 50 Mbp (a quarter of the chromosome). The ASGR includes highly repetitive sequences from an Opie-2-like retrotransposon family that are particularly abundant in this region of the genome.     

Apomixis in agriculture: the quest for clonal seeds

Apomixis, or asexual reproduction through seeds, is a natural trait that could have an immense positive impact on crop production. Apomictic breeding strategies could allow the fixation and indefinite propagation of any desired genotype, however complex. Apomicts display a wide variety of developmental mechanisms, which can be viewed as a short-circuiting of sexual development. Gametophytic and sporophytic apomixis are distinguished by the developmental origin of apomictically derived embryos. Genetic studies suggest that individual elements of gametophytic apomixis, such as apomeiosis and parthenogenesis, are either controlled by one or two dominant Mendelian factors. As recombination around apomeiosis loci is suppressed, it is currently not known how complex these loci are. Much less is known regarding the genetic control of sporophytic apomixis but initial studies suggest a complex genetic control. Genetic analyses of sexual reproduction in plant model systems have identified genes that, when mutated, display elements of apomixis. Such studies help in the identification of candidate genes and promoters that can be used for the de novo engineering of apomixis through biotechnology. Molecular genetic studies in apomictic and sexual systems will generate the knowledge necessary for the engineering of conditional apomixis technology. Approaches encouraging collaboration and widespread dissemination of the acquired knowledge will constitute the most innovative route to the development, deployment and acceptance of apomixis technology in agriculture.     

Differential effects of polyploidy and diploidy on fitness of apomictic Boechera

The co-occurrence of apomixis (asexual reproduction) and polyploidy in plants has been the subject of debate in regard to the origin and evolution of asexuality. In recent years, polyploidy has been postulated as a maintenance and stabilization factor rather than as a source of apomixis origin. It is assumed polyploidy facilitates the compensation for mutation accumulation, and hence, the rare occurrence of diploid apomixis indirectly supports this finding. Nevertheless, diploid apomicts exist and are successful, especially in the genus Boechera. While comparing phenotypic traits, fitness-related traits and apomixis penetrance between both diploid and triploid apomicts in the genus Boechera, it was expected to find trait variance that can be attributed to ploidy. Surprisingly, little trait variation could be assigned to ploidy, but rather trait variations were mainly genotype-specific. Additionally, it is shown that paternal contribution is very important for trait success, even though all offspring are genetically identical to the mother plant. This harbors implications for the introduction of apomixis into crop plants, considering the effects of paternal contribution during asexual reproduction. Nevertheless, polyploidy is an efficient way to buffer deleterious mutations, but the flexibility of diploid apomicts of the genus Boechera for rare sexual events contributes to their success in nature.     

The Genetic Control of Apomixis: Asexual Seed Formation

Apomixis (asexual seed formation) is the result of a plant gaining the ability to bypass the most fundamental aspects of sexual reproduction: meiosis and fertilization. Without the need for male fertilization, the resulting seed germinates a plant that develops as a maternal clone. This dramatic shift in reproductive process has been documented in many flowering plant species, although no major seed crops have been shown to be capable of apomixis. The ability to generate maternal clones and therefore rapidly fix desirable genotypes in crop species could accelerate agricultural breeding strategies. The potential of apomixis as a next-generation breeding technology has contributed to increasing interest in the mechanisms controlling apomixis. In this review, we discuss the progress made toward understanding the genetic and molecular control of apomixis. Research is currently focused on two fronts. One aims to identify and characterize genes causing apomixis in apomictic species that have been developed as model species. The other aims to engineer or switch the sexual seed formation pathway in non-apomictic species, to one that mimics apomixis. Here we describe the major apomictic mechanisms and update knowledge concerning the loci that control them, in addition to presenting candidate genes that may be used as tools for switching the sexual pathway to an apomictic mode of reproduction in crops.