拟南芥成花关键基因调控网络研究进展

开花是植物从营养生长转变为生殖生长的重要时期,而开花调控成为近年来植物分子生物学研究的热点。在目前已有的研究中,调控拟南芥开花的基因网络已经发展成一个包含串扰(Crosstalk)、反馈(Feedback)和冗余(Redundancy)的复杂网络,这个网络通过开花整合子来与其他发育过程紧密结合。以调节开花的遗传途径作为基础,重点讨论了顶端分生组织中的信号积累、花发育的时空调节、开花相关基因在拟南芥开花时间或花发育过程以外的其他过程中的功能,并对开花调控网络的深入研究进行了展望。     

LEAFY同源基因研究进展

LEAFY(LFY)同源基因存在于所有的陆生植物中,在植物花发育早期表达,并在花发育过程中抑制茎端分生组织的营养生长,调控花分生组织和花器官的形成,使转LFY基因植株提前开花,LFY同源基因与其上下游基因共同调控花发育过程.LFY同源基因的蛋白质结构在不同物种间保守性很高,但它们的表达部位差异很大.该文总结了近年来国内外已经克隆到的LFY同源基因的表达、功能及其在果树、花卉、粮食作物上的应用,以期为植物花发育的深入研究提供参考.     

拟南芥春化作用相关基因FLC的表达调控及天然早花突变体的研究进展

植物开花是从营养生长到生殖状态的重要发育转变,是多种内在因子和环境因素共同作用的结果。在拟南芥开花调控网络中,开花抑制基因FLC处于枢纽地位。FLC的表达受许多来自环境和生长发育的信号调控,主要包括：PAF1复合体、SWR1复合体成员,FRI依赖途径、自主途径和春化作用途径基因。本文主要综述了影响FLC表达的春化相关基因及天然早花突变体的研究进展,并根据最新的研究成果提出该研究领域的研究方向和重点。     

花序分生组织特性基因TFL1的系统发育及其功能分析

TFL1同源基因在维持植物营养生长和花序分生组织特性方面起着非常重要的作用,其功能的丧失常导致植物提早开花,花序的正常发育受到抑制,最终茎端形成顶花。至今已经有28种植物的TFL1基因被克隆到,其中包括拟南芥、金鱼草和番茄等模式植物。TFL1 蛋白的系统发育树基本符合物种的亲缘关系。作为花序分生组织特性基因的TFL1与花分生组织特性基因LFY 和AP1相互作用,抑制花序分生组织向花分生组织的转变。TFL1和LFY等基因可用来培育早花新品种,也可用于培育无果的新品种,减少悬铃木、杨、柳等果毛的污染。     

拟南芥花分生组织决定基因AGL24对花发育的影响

AGAMOUS-LIKE 24(AGL24)基因编码MADS蛋白,在植物花发育的不同时期发挥着重要的作用。综述了AGL24如何通过和其他花分生组织决定基因的相互作用来影响拟南芥花的发育,调节开花时间,这将有助于人们对开花基因调控网络有更进一步的认识,能够在生产上有效的调控开花时间,从而为植物育种提供借鉴。     

成花素FT的调控运输及生理功能的研究进展

开花作为植物从营养生长到生殖生长的转折点,受外界环境变化影响,对植物发育和繁殖有着至关重要的作用。成花素(flowering locus T FT)蛋白作为拟南芥开花信号核心分子,在开花调控中扮演重要角色。拟南芥叶片在感应外界温度光照等因素变化后,通过生理钟(constants,CO)等蛋白及部分Mico-RNA正负协调调控筛管伴胞中的FT基因表达,FT蛋白通过筛管从叶片运输到茎顶端分生组织后,与一个碱性亮氨酸拉链(basic leucine zipper,b ZIP)蛋白FD结合调控茎顶端分生组织(shoot apical meristem,SAM)中的花组织形成相关基因表达,继而诱导开花。对近年来FT基因在叶片中的调控、FT蛋白的运输及其在顶端分生组织中的开花诱导机理进行综述,为进一步完善FT表达调控及功能研究提供参考。     

植物开花的组蛋白甲基化调控分子机理

开花是指植物从营养生长转变到生殖生长的生理过程, 是植物个体发育和后代繁衍的中心环节, 既受遗传基础决定,同时又受到温度和光周期等多种环境因素的调控。在拟南芥中, 已经分离了大量的与开花相关的基因, 从遗传学上已初步形成了一个开花调控的网络。组蛋白甲基化是植物发育过程的重要调节方式, 近年来关于其参与开花调控的研究有了重要进展。本文综述了具有代表性的组蛋白H3赖氨酸甲基化修饰参与调控植物开花发育的机制, 提出该研究领域的发展方向和前景。     

植物开花的组蛋白甲基化调控分子机理

开花是指植物从营养生长转变到生殖生长的生理过程,是植物个体发育和后代繁衍的中心环节,既受遗传基础决定,同时又受到温度和光周期等多种环境因素的调控。在拟南芥中,已经分离了大量的与开花相关的基因,从遗传学上已初步形成了一个开花调控的网络。组蛋白甲基化是植物发育过程的重要调节方式,近年来关于其参与开花调控的研究有了重要进展。本文综述了具有代表性的组蛋白H3赖氨酸甲基化修饰参与调控植物开花发育的机制,提出该研究领域的发展方向和前景。     

植物MADS-box 基因FRUITFULL(FUL)研究进展

植物MADS-box基因家族的不同成员在植物生长发育过程中起着非常重要的作用。拟南芥MADS-box 基因FRUITFULL(FUL) 在控制拟南芥开花时间、花分生组织分化、茎生叶形态以及心皮和果实的发育中起到重要作用。其他植物中,FUL的同源基因也在调控花发育,果实发育以及叶片发育等方面各自起到重要作用。本文综述了FUL基因及其同源基因的表达模式和功能,并就其在农作物及果树育种上的潜在应用价值进行了讨论。     

日本晚樱PrseSHP基因的克隆与功能分析

采用同源克隆方法结合RACE技术,从日本晚樱(Prunus lannesiana)品种‘大岛樱’中克隆到花发育调控相关的PrseSHP基因(GenBank登录号为GU362645)。PrseSHP基因序列全长1 223bp,包含1个长741bp的完整开放阅读框,编码246个氨基酸和1个终止密码子。分子系统发生分析表明,PrseSHP属MADS-box转录因子的PLE/SHP进化系,并与蔷薇科植物的SHP同源蛋白聚于同一进化分支;蛋白序列比对显示,该转录因子拥有M、I、K和C共4个结构域,且其C末端结构域中包含高度保守的AGⅠ和Ⅱ基序。基因表达分析表明,PrseSHP基因主要在‘大岛樱’的花瓣、雄蕊、雌蕊和幼果等器官中表达,在花萼中仅能检测到微弱的转录信号,在幼叶中不表达,与其他植物SHP同源基因的表达模式有一定的差别。功能分析显示,转PrseSHP基因拟南芥植株明显比野生型拟南芥弱小,转基因拟南芥在6~8片莲座叶后即抽薹开花,时间较野生型拟南芥(14~17片莲座叶后抽薹开花)明显提前,证明异位表达的PrseSHP基因能促进拟南芥早花,其在花发育过程中可能参与调控植物开花。     

拟南芥LEAFY基因在花发育中的网络调控及其生物学功能

重点综述了拟南芥花分生组织特征基因——LEAFY（LFY）基因及其同源基因在花发育中的网络调控及其生物学功能。LFY基因广泛表达于高等植物的营养性和生殖性组织。LFY基因需要与其他基因相互作用,並且表达量达到一定水平时才能促进成花。LFY基因处于成花调控网络的关键位置,不仅调控开花时间和花转变,而且在花序和花的发育中也起重要作用。碳源、植物激素等因子直接或间接地影响LFY基因的表达和作用。提示通过掌握LFY基因的表达调控规律进一步探讨成花机理的可行性。 Abstract:Recent research progress on regulation network and biological roles of LFY gene in Arabidopsis thaliana and its homologue genes in floral development are reviewed emphatically in the present paper.LFY gene expresses widely in both vegetative and reproductive tissues in different higher plants,therefore investigation on role of LFY gene on flowering is of general significance.LFY gene plays an important role to promote flower formation by interaction and coordination with other genes,such as TFL,EMF,AP1,AP2,CAL,FWA,FT,AP3,PI,AG,UFO,CO,LD,GA1 etc,and a critical level of LFY expression is essential.LFY gene not only controls flowering-time and floral transition,but also plays an important role in inflorescence and floral organ development.It was situated at the central site in gene network of flowering regulation,positively or negatively regulates the level or activities of flowering-related genes.Some physiological factors,such as carbon sources,phytohormones,affect directly or indirectly the expression and actions of LFY gene.This indicates that level of LFY expression can also be regulated with physiological methods.It is probable that we can explain the principal mechanism of flowering by regulation network of LFY gene.     

EMF1 interacts with EIP1, EIP6 or EIP9 involved in the regulation of flowering time in Arabidopsis

The EMBRYONIC FLOWER (EMF) 1 gene has been shown to be necessary for maintenance of vegetative development. To investigate the molecular mechanism of EMF1-mediated plant development, we screened EMF1-interacting proteins and identified 11 candidate proteins using the yeast two-hybrid system. Among the candidate genes, three EMF1-Interacting Protein (EIP) genes, EIP1, EIP6 and EIP9, are predicted to encode a WNK (with-no-lysine) kinase, a B-box zinc-finger protein and a DnaJ-domain protein, respectively. The expression patterns of EIP1, EIP6 and EIP9 were similar to that of EMF1, and EMF1-EIP1, EMF1-EIP6 and EMF1-EIP9 heterodimers were localized in the nucleus. In addition, eip1, eip6 and eip9 mutants flowered early and showed increased expression of flowering-time and floral organ identity genes, while EIP1-, EIP6- and EIP9-overexpressing transgenic plants showed late flowering phenotypes. Our results suggest that EMF1 interacts with EIP1, EIP6 and EIP9 during vegetative development to regulate flowering time in Arabidopsis.     

Mechanisms and function of flower and inflorescence reversion

Flower and inflorescence reversion involve a switch from floral development back to vegetative development, thus rendering flowering a phase in an ongoing growth pattern rather than a terminal act of the meristem. Although it can be considered an unusual event, reversion raises questions about the nature and function of flowering. It is linked to environmental conditions and is most often a response to conditions opposite to those that induce flowering. Research on molecular genetic mechanisms underlying plant development over the last 15 years has pinpointed some of the key genes involved in the transition to flowering and flower development. Such investigations have also uncovered mutations which reduce floral maintenance or alter the balance between vegetative and floral features of the plant. How this information contributes to an understanding of floral reversion is assessed here. One issue that arises is whether floral commitment (defined as the ability to continue flowering when inductive conditions no longer exist) is a developmental switch affecting the whole plant or is a mechanism which assigns autonomy to individual meristems. A related question is whether floral or vegetative development is the underlying default pathway of the plant. This review begins by considering how studies of flowering in Arabidopsis thaliana have aided understanding of mechanisms of floral maintenance. Arabidopsis has not been found to revert to leaf production in any of the conditions or genetic backgrounds analysed to date. A clear-cut reversion to leaf production has, however, been described in Impatiens balsamina. It is proposed that a single gene controls whether Impatiens reverts or can maintain flowering when inductive conditions are removed, and it is inferred that this gene functions to control the synthesis or transport of a leaf-generated signal. But it is also argued that the susceptibility of Impatiens to reversion is a consequence of the meristem-based mechanisms controlling development of the flower in this species. Thus, in Impatiens, a leaf-derived signal is critical for completion of flowering and can be considered to be the basis of a plant-wide floral commitment that is achieved without accompanying meristem autonomy. The evidence, derived from in vitro and other studies, that similar mechanisms operate in other species is assessed. It is concluded that most species (including Arabidopsis) are less prone to reversion because signals from the leaf are less ephemeral, and the pathways driving flower development have a high level of redundancy that generates meristem autonomy even when leaf-derived signals are weak. This gives stability to the flowering process, even where its initiation is dependent on environmental cues. On this interpretation, Impatiens reversion appears as an anomaly resulting from an unusual combination of leaf signalling and meristem regulation. Nevertheless, it is shown that the ability to revert can serve a function in the life history strategy (perenniality) or reproductive habit (pseudovivipary) of many plants. In these instances reversion has been assimilated into regular plant development and plays a crucial role there.     

Genetics of flower initiation and development in annual and perennial plants

Flowering is an integral developmental process in angiosperms, crucial to reproductive success and continuity of the species through time. Some angiosperms complete their life cycle within a year (annual plants), and others have a longer reproductive life, which is characterized by the generation of new flowering and vegetative shoots every year (perennial plants). Despite the differences in their lifespan, the underlying genetics of flower induction and floral organ formation appears to be similar among these plants. Hence, the knowledge gained from the study of flowering mechanism in Arabidopsis thaliana can be used to better understand similar processes in other plant species, especially the perennials, which usually have a long generation time and are not amenable to genetic analysis. Using Arabidopsis as a model, we briefly discuss the current understanding of the transition from vegetative to reproductive growth and the subsequent formation of individual floral organs, and how this knowledge has been successfully applied to the identification of homologous genes from perennial crops. Although annuals appear to share many similarities with perennials in terms of gene function, they differ in their commitment to flowering. Once an annual reaches the reproductive phase, all meristems are typically converted into either floral or inflorescence meristems. In contrast, each year, each meristem of a mature perennial has the choice to produce either a vegetative or a reproductive shoot. The physiology and genetics of flowering in Citrus are used to highlight the complexity of reproductive development in perennials, and to discus possible future research directions.     

early in short days 4, a mutation in Arabidopsis that causes early flowering and reduces the mRNA abundance of the floral repressor FLC

The plant shoot is derived from the apical meristem, a group of stem cells formed during embryogenesis. Lateral organs form on the shoot of an adult plant from primordia that arise on the flanks of the shoot apical meristem. Environmental stimuli such as light, temperature and nutrient availability often influence the shape and identity of the organs that develop from these primordia. In particular, the transition from forming vegetative lateral organs to producing flowers often occurs in response to environmental cues. This transition requires increased expression in primordia of genes that confer floral identity, such as the Arabidopsis gene LEAFY. We describe a novel mutant, early in short days 4 (esd4), that dramatically accelerates the transition from vegetative growth to flowering in Arabidopsis: The effect of the mutation is strongest under short photoperiods, which delay flowering of Arabidopsis: The mutant has additional phenotypes, including premature termination of the shoot and an alteration of phyllotaxy along the stem, suggesting that ESD4 has a broader role in plant development. Genetic analysis indicates that ESD4 is most closely associated with the autonomous floral promotion pathway, one of the well-characterized pathways proposed to promote flowering of Arabidopsis: Furthermore, mRNA levels of a floral repressor (FLC), which acts within this pathway, are reduced by esd4, and the expression of flowering-time genes repressed by FLC is increased in the presence of the esd4 mutation. Although the reduction in FLC mRNA abundance is likely to contribute to the esd4 phenotype, our data suggest that esd4 also promotes flowering independently of FLC. The role of ESD4 in the regulation of flowering is discussed with reference to current models on the regulation of flowering in Arabidopsis.     

Regulation of floral initiation in horticultural trees

The intention of this review is to discuss floral initiation of horticultural trees. Floral initiation is best understood for herbaceous species, especially at the molecular level, so a brief overview of the control of floral initiation of Arabidopsis (Arabidopsis thaliana (L.) Heynh.) precedes the discussion of trees. Four major pathways to flowering have been characterized in Arabidopsis, including environmental induction through photoperiod and temperature, autonomous floral initiation, and regulation by gibberellins. Tropical trees are generally induced to flower through environmental cues, whereas floral initiation of temperate deciduous trees is often autonomous. In the tropical evergreen tree mango, Mangifera indica L., cool temperature is the only factor known to induce flowering, but does not ensure floral initiation will occur because there are important interactions with vegetative growth. The temperate deciduous tree apple, Malus domestica Borkh., flowers autonomously, with floral initiation dependent on aspects of vegetative development in the growing season before anthesis, although with respect to the floral initiation of trees in general: the effect of the environment, interactions with vegetative growth, the roles of plant growth regulators and carbohydrates, and recent advances in molecular biology, are discussed.     

Floral organ identity genes in the orchid Dendrobium crumenatum

Orchids are members of Orchidaceae, one of the largest families in the flowering plants. Among the angiosperms, orchids are unique in their floral patterning, particularly in floral structures and organ identity. The ABCDE model was proposed as a general model to explain flower development in diverse plant groups, however the extent to which this model is applicable to orchids is still unknown. To investigate the regulatory mechanisms underlying orchid flower development, we isolated candidates for A, B, C, D and E function genes from Dendrobium crumenatum. These include AP2-, PI/GLO-, AP3/DEF-, AG- and SEP-like genes. The expression profiles of these genes exhibited different patterns from their Arabidopsis orthologs in floral patterning. Functional studies showed that DcOPI and DcOAG1 could replace the functions of PI and AG in Arabidopsis, respectively. By using chimeric repressor silencing technology, DcOAP3A was found to be another putative B function gene. Yeast two-hybrid analysis demonstrated that DcOAP3A/B and DcOPI could form heterodimers. These heterodimers could further interact with DcOSEP to form higher protein complexes, similar to their orthologs in eudicots. Our findings suggested that there is partial conservation in the B and C function genes between Arabidopsis and orchid. However, gene duplication might have led to the divergence in gene expression and regulation, possibly followed by functional divergence, resulting in the unique floral ontogeny in orchids.