多倍体植物的表观遗传现象

表观遗传现象是指基因表达发生改变但不涉及DNA序列的变化, 它存在于许多植物的多倍体化过程中,而且能够在代与代之间传递。表观遗传变异包括基因沉默、DNA甲基化、核仁显性、休眠转座子激活和基因组印记等方面。这种现象可能是由于基因组间的相互作用直接诱发基因沉默或基因表达改变所致;也可能由DNA甲基化之外的组蛋白编码的改变引起;或者与甲基化不足、染色质重组或转座子激活等有关。表观遗传变异在提高基因表达的多样性,引起遗传学和细胞学上的二倍化,以及促进基因组间的相互协调等方面起着重要作用。文章综述了植物多倍体化过程中的表观遗传现象及其在多倍体植物基因组进化中的作用,并在此基础上提出了今后在这方面的研究途径。     

植物多倍体化中基因组和基因表达的变化

多倍体化在植物进化的历史过程中频繁发生, 对新物种的形成产生了很大影响。伴随着多倍体化, 植物在基因组和基因表达上发生了复杂的变化, 包括染色体数目变化、染色体重组、基因沉默、基因的非加性表达和表观遗传等变化。该文对多倍体化引起的这些变化及其相应的机理进行了综述, 以期为了解多倍体化中植物新表型的产生机理和在进化中的意义提供参考。     

植物多倍体研究的回顾与展望

多倍化是促进植物进化的重要力量。多倍体主要是通过未减数配子融合,体细胞染色体加倍以及多精受精三种方式起源的。其中,不减数配子是多倍体形成的主要机制。三倍体可能在四倍体的进化中起了重要作用。过去认为多倍体只能是进化的死胡同,现在发现很多多倍体类群都是多元起源的而不是单元起源的。当多倍体形成后,基因组中的重复基因大部分保持原有的功能,也有相当比例的基因发生基因沉默。多倍体通常表现出不存在于二倍体祖先的表型,并且超出了其祖先的分布范围,因为在多倍体中发生了很多基因表达的变化。主要从多倍体的起源、影响多倍体发生的因素及多倍体基因组的进化等方面回顾并展望多倍体的研究。     

多倍体植物混合倍性种群的建立机制研究进展

基因组多倍化是物种形成和进化的重要驱动力, 几乎所有植物都经历过至少一次基因组加倍。然而, 由于多倍体植株比二倍体表现出更高的死亡率, 多倍化机制被认为是植物进化的“死胡同”。一些植物物种具有自然混合倍性种群, 即同一物种具有不同倍性, 这为揭示多倍体的进化机制提供了最佳途径。本文从基因组加倍形成多倍体植物开始, 综述了混合倍性种群的形成、建立与维持的研究进展, 探讨了多倍体适应自然环境的种群分化而形成多倍体物种的机制。研究自然混合倍性种群的倍性组成、重复基因的功能分化以及多倍体的生态位分化, 有利于明确混合倍性自然种群的生态适应与维持机理, 以及多倍体植物的进化机制。     

荧光原位杂交技术在植物多倍体起源与进化研究中的应用

多倍化是植物物种形成与多样化的重要原动力。研究植物特别是一些重要经济作物和园艺植物多倍体的起源与进化,不仅对于揭示多倍体形成过程中性状变异的分子机制具有重要意义,而且可为植物遗传资源的保护与利用提供理论和技术支持。作为连接基因组序列片段到染色体组的桥梁,荧光原位杂交技术长期被广泛用来研究多倍体形成与进化过程中相关特异基因或序列的表达定位、外源染色体检测和鉴定、基因组结构变异等科学问题。因此,在简单介绍荧光原位杂交技术发展历史和植物多倍体主要类型的基础上,主要总结了荧光原位杂交技术在植物多倍体起源与进化相关研究上的应用。     

非编码RNA与哺乳动物基因组印记的起源

基因组印记是由亲本来源不同而导致等位基因表达差异的一种遗传现象,主要发生在胎盘哺乳动物（真哺乳类）和显花植物中.大部分印记基因都分布在印记基因簇内,其中包含大量的非编码RNA基因.印记基因的表达受印记控制区（ICRs）的顺式调控.基因组印记产生的原因及过程是现代遗传学研究的一个热点问题,分析印记同源区从非印记物种到印记物种的过渡,为解决这一问题提供了重要启示.最近,原始哺乳动物（有袋类和单孔类）模式物种全基因组测序的完成,极大地促进了印记同源区的比较分析研究.本文对这些研究进行了回顾和分析,发现非编码RNA与哺乳动物基因组印记获得关系密切.主要依据为：（1）伴随着基因组印记的获得,印记区有大量的非编码RNA新基因出现;（2）与基因组印记相关的一些保守非编码RNA的表达发生了显著变化.此外,对15种脊椎动物中印记snoRNA基因系统分析的结果表明：印记snoRNA起源于真哺乳类与有袋类动物分化之后,并且在真哺乳类辐射进化之前发生了迅速的扩张,主要的基因家族在这一时期已经形成.这些结果进一步证明了非编码RNA与基因组印记获得的密切联系.非编码RNA可能主要通过调控印记表达和诱导染色体表观遗传修饰两种机制,参与哺乳动物基因组印记的获得.     

植物人工异源多倍体的遗传及后遗传变化

据估计,70%以上的显花植物在其生活史上至少发生过一次以上的多倍化。传统的有关多倍性的观点认为,多倍体基因组应是其双亲基因组的积加。但是,有些合成异源多倍体的基因组发生了广泛的遗传及后遗传变化。这些变化包括亲本DNA序列丢失、核仁显性、DNA甲基化模式改变、基因沉默、反转座子激活等。亲本序列丢失可能与部分同源序列间重组有关,而亲本基因沉默可能与同源性依赖的基因沉默及RNA干涉等有关。     

植物多倍化与多倍体基因组进化研究进展

多倍化(polyploidization)是指细胞核中的染色体组发生加倍并以可遗传的方式传递至后代的现象.虽然已有研究揭示多倍化事件普遍出现于被子植物各类群的进化过程中,但其对物种多样化与基因组进化的作用始终都处于争论之中.近年来随着基因组测序的革命性进步与多种组学和分子生物学技术的应用,植物多倍化与多倍体基因组进化领域的研究已取得多方面的重要进展.本文首先系统地介绍了植物多倍化的研究历史、多倍体分类系统以及该领域目前存在的主要学术争论.在此基础上,侧重从染色体数目与结构、DNA和组蛋白表观遗传修饰以及RNA和蛋白质表达等多个层次,对在多倍体小麦、油菜与棉花等模式作物中所取得的研究成果进行了较详细的概括.期望本文通过对最新研究成果的总结与未来研究展望,进一步增进对多倍化在植物物种多样性形成与基因组进化过程中重要作用的理解,促进我国植物多倍化研究领域的发展.     

母源效应蛋白在基因组印记维持中的作用

基因组印记是指生殖细胞发生过程中双亲基因组发生差异表观修饰,使带有亲代印记的等位基因出现父源或母源单等位基因表达。在配子发生和早期胚胎发育过程中,基因组印记甲基化经历一个去除、重建和维持的复杂过程。这个过程中的任何环节被干扰都将导致印记紊乱,造成胚胎发生、胎盘形成及出生后发育异常。近来研究表明,早期胚胎发育过程中一些母源效应蛋白在印记基因表观调控中起重要作用。为了更好地理解这些母源因子对印记基因建立及维持的作用与机制,文章综述了DPPA3、ZFP57、TRIM28和DNMT1等母源效应因子近年来的相关研究进展,并探讨了这些因子对基因组印记的表观调控机制。     

植物同源多倍体耐盐性研究进展

多倍化是高等植物进化最重要的动力之一,多倍体植物由于基因组组成以及基因表达方面的变化,通常会表现出不同的生理现象,多倍体的抗性优于其同源二倍体祖先。土壤盐碱化和次生盐渍化是影响农作物生产的重要因素,严重制约着我国农业的可持续发展。同源多倍体植物耐盐能力较强,是作物遗传改良的重要种质资源,了解其耐盐机理对培育耐盐品种具有重要意义。本文从与盐胁迫相关的耐盐性进化、生理生化水平、细胞结构和分子层面等多角度总结了植物同源多倍体盐胁迫研究进展,并以作者所在研究团队培育出的多倍体西瓜为例讨论了多倍体抗逆性研究存在的问题及未来的发展方向,以期为多倍体抗逆优势机理研究提供参考。     

Genome evolution in Oryza allopolyploids of various ages: Insights into the process of diploidization

The prevalence and recurrence of whole-genome duplication in plants and its major role in evolution have been well recognized. Despite great efforts, many aspects of genome evolution, particularly the temporal progression of genomic responses to allopolyploidy and the underlying mechanisms, remain poorly understood. The rice genus Oryza consists of both recently formed and older allopolyploid species, representing an attractive system for studying the genome evolution after allopolyploidy. In this study, through screening BAC libraries and sequencing and annotating the targeted BAC clones, we generated orthologous genomic sequences surrounding the DEP1 locus, a major grain yield QTL in cultivated rice, from four Oryza polyploids of various ages and their likely diploid genome donors or close relatives. Based on sequenced DEP1 region and published data from three other genomic regions, we investigated the temporal evolutionary dynamics of four polyploid genomes at both genetic and expression levels. In the recently formed BBCC polyploid, Oryza minuta, genome dominance was not observed and its short-term responses to allopolyploidy are mainly manifested as a high proportion of homoeologous gene pairs showing unequal expression. This could partly be explained by parental legacy, rewiring of divergent regulatory networks and epigenetic modulation. Moreover, we detected an ongoing diploidization process in this genus, and suggest that the expression divergence driven by changes of selective constraint probably plays a big role in the long-term diploidization. These findings add novel insights into our understanding of genome evolution after allopolyploidy, and could facilitate crop improvements through hybridization and polyploidization.     

Genome evolution in polyploids

Polyploidy is a prominent process in plants and has been significant in the evolutionary history of vertebrates and other eukaryotes. In plants, interdisciplinary approaches combining phylogenetic and molecular genetic perspectives have enhanced our awareness of the myriad genetic interactions made possible by polyploidy. Here, processes and mechanisms of gene and genome evolution in polyploids are reviewed. Genes duplicated by polyploidy may retain their original or similar function, undergo diversification in protein function or regulation, or one copy may become silenced through mutational or epigenetic means. Duplicated genes also may interact through inter-locus recombination, gene conversion, or concerted evolution. Recent experiments have illuminated important processes in polyploids that operate above the organizational level of duplicated genes. These include inter-genomic chromosomal exchanges, saltational, non-Mendelian genomic evolution in nascent polyploids, inter-genomic invasion, and cytonuclear stabilization. Notwithstanding many recent insights, much remains to be learned about many aspects of polyploid evolution, including: the role of transposable elements in structural and regulatory gene evolution; processes and significance of epigenetic silencing; underlying controls of chromosome pairing; mechanisms and functional significance of rapid genome changes; cytonuclear accommodation; and coordination of regulatory factors contributed by two, sometimes divergent progenitor genomes. Continued application of molecular genetic approaches to questions of polyploid genome evolution holds promise for producing lasting insight into processes by which novel genotypes are generated and ultimately into how polyploidy facilitates evolution and adaptation.     

Non-coding RNAs, epigenetics and complexity

Several aspects of epigenetics are strongly linked to non-coding RNAs, especially small RNAs that can direct the cytosine methylation and histone modifications that are implicated in gene expression regulation in complex organisms. A fundamental characteristic of epigenetics is that the same genome can show alternative phenotypes, which are based in different epigenetic states. Some of the most studied complex epigenetic phenomena including transposon activity and silencing recently exemplified by piRNAs (piwi-interacting RNAs), position effect variegation, X-chromosome inactivation, parental imprinting, and paramutation have direct or indirect participation of an RNA component. Conceivably, most of the non-coding RNAs with no described function yet, are players in epigenetic mechanisms that are still not completely understood. In that regard, RNAs were recently implicated in new mechanisms of genetic information transfer in yeast, plants and mice. In this review article, the hypothesis that non-coding RNAs might be the main component of complex organisms acquired during evolution will be explored. The question of how evolutionary theories have been challenged by these molecules in association with epigenetic mechanisms will also be discussed here.     

植物多倍体基因组的形成与进化

多倍化是植物进化变异的自然现象,也是促进植物发生进化改变的重要力量。在被子植物中,约 70％的种类在进化史中曾发生过一次或多次多倍化的过程。目前的研究结果表明,自然界绝大多数多倍体是通过未减数配子的融合而形成的,并且很多多倍体种是通过多次独立的多倍化过程而重复发生的。由多倍化所导致的重复基因在多倍体基因组中可能有三种不同的命运,即：保持原有的功能、基因沉默或分化并执行新的功能。多倍化以后,重复基因组的进化动态则主要表现在染色体重排和“染色体二倍化”、不同基因组之间的相互渗透、以及核-质之间的相互作用等方面。     

Size matters in Triticeae polyploids: larger genomes have higher remodeling

Polyploidization is one of the major driving forces in plant evolution and is extremely relevant to speciation and diversity creation. Polyploidization leads to a myriad of genetic and epigenetic alterations that ultimately generate plants and species with increased genome plasticity. Polyploids are the result of the fusion of two or more genomes into the same nucleus and can be classified as allopolyploids (different genomes) or autopolyploids (same genome). Triticeae synthetic allopolyploid species are excellent models to study polyploids evolution, particularly the wheat-rye hybrid triticale, which includes various ploidy levels and genome combinations. In this review, we reanalyze data concerning genomic analysis of octoploid and hexaploid triticale and different synthetic wheat hybrids, in comparison with other polyploid species. This analysis reveals high levels of genomic restructuring events in triticale and wheat hybrids, namely major parental band disappearance and the appearance of novel bands. Furthermore, the data shows that restructuring depends on parental genomes, ploidy level, and sequence type (repetitive, low copy, and (or) coding); is markedly different after wide hybridization or genome doubling; and affects preferentially the larger parental genome. The shared role of genetic and epigenetic modifications in parental genome size homogenization, diploidization establishment, and stabilization of polyploid species is discussed.     

Parental genomic imprinting in plants: significance for reproduction

Parental genomic imprinting is an epigenetic phenomenon causing the expression of a gene from one of the two parental alleles. Imprinting has been identified in plants and mammals. Recent evidence shows that DNA methylation and histone modifications are responsible for this parent-of-origin dependent expression of imprinted genes. We review the mechanisms and functions of imprinting in plants. We further describe the significance of imprinting for reproduction and discuss potential models for its evolution.