被子植物起源和早期演化研究的回顾与展望

近年来,被子植物起源和早期演化研究,由于手段和技术的更新,资料大量积累,取得了许多重要进展,成为植物学领域的一大热点。本文对过去近五十年的研究作了回顾,并从分子系统学、分支系统学、花原基发生的形态学、花发育的分子遗传学及白垩纪花和其它生殖结构化石研究等五个方面对该领域在最近十几年的研究进展进行综述,最后,对今后如何开展这方面的工作作了简要评论。     

花对称性的研究进展

花对称性(floral symmetry)是被子植物花部结构的典型特性之一,主要有辐射对称和两侧对称两种形式。被子植物初始起源的花为辐射对称,而两侧对称的花则是由辐射对称的花演变而来。两侧对称的花部结构是被子植物进化过程中的一个关键的革新,被认为是物种形成和分化的关键推动力之一。近年来有关花对称性的形成和进化机制的研究在植物学科的不同领域均取得了长足的进展。本文综述了花对称性在发育生物学、传粉生物学、生殖生态学及分子生物学等方面的研究进展。两侧对称形成于被子植物花器官发育的起始阶段,随后贯穿整个花器官发育过程或者出现在花器官发育后期的不同阶段。花器官发育过程中一种或多种类型器官的败育以及特异性花器官结构的形成是两侧对称形成的主要原因。研究表明,在传粉过程的不同阶段,花对称性均会受到传粉昆虫介导的选择作用。相比辐射对称的花,两侧对称的花提高了特异性传粉者的选择作用,增加了花粉落置的精确性,进而确保了其生殖成功。花对称性的分子机理已经在多种双子叶植物中进行了深入的研究。现有的证据表明,CYC同源基因在花对称性的分子调控方面起着非常重要的作用。花对称性在被子植物进化过程中是如何起源,与其他花部构成之间是否协同作用,一些不符合一般模式的科属其花对称性的形成机制等都是今后要进一步研究的命题。     

被子植物花器官发育的模型演变和分子调控

基于模式植物拟南芥(Arabidopsis thaliana)和金鱼草(Antirrhinum majus)花器官突变体研究提出的四聚体模型揭示了花同源异型蛋白的相互作用方式;进一步提出的核小体拟态模型,解释了花同源蛋白四聚体调控目标靶基因的分子机理。被子植物花器官形态多样化与MADS-box基因的表达模式和功能分化密切相关。多年生被子植物花发育的高通量转录组分析表明,多种基因参与调控花器官发育过程。本文重点综述了被子植物花器官发育的模型演变、MADS-box基因结构和基因重复、miRNA调控以及相关转录组分析的最新研究成果,并对花器官发育的研究前景进行了展望。     

Pollinator-dependent evolution of floral trait combinations in an orchid herb

作为驱动花性状进化的重要媒介之一,传粉者塑造了被子植物多样的花部特征。目前,大部分研究集中于传粉者驱动被子植物单个性状的进化,而较少涉及对传粉者驱动被子植物组合性状进化的研究。本研究以兰科植物绶草(Spiranthes sinensis)为材料,分别鉴别和估算了传粉者对植物单个性状和组合性状施加的定向选择和关联选择压力。研究结果表明,传粉者对开花时间和花大小施加定向选择压力,传粉者选择早开花、花更大的个体。传粉者对绶草花展示的组合性状(花大小与花数目)施加正向的关联选择压力,而对开花时间与花展示的组合性状(开花时间与株高、开花时间与花数目)施加负向的关联选择压力。同时,传粉者的选择强度在年际间存在差异。该研究证实了传粉者在驱动植物组合性状进化过程中的作用,同时强调在认识和理解被子植物表型性状进化轨迹时,需要考虑植物组合性状的影响。     

生花植物概念简介及其名称的商榷

在上个世纪最后的 2 0年里 ,系统学家应用形态性状对种子植物进行了大量的分支分析。其结果显示灭绝的五柱木属加上灭绝的本内苏铁目及尚存的买麻藤目是被子植物的姊妹群 ,形成一个强支 ,称之为生花植物支。生花植物假说对探讨被子植物起源有着重要影响 ,它激发人们讨论被子植物起源时间可能要提前到三叠纪甚至石炭纪 ,除了支持原有的真花学说外 ,还提出新恩格勒学说。但是 ,近年来对现存种子植物进行分子系统学研究的结果是 :(1)拒绝接受生花植物概念 ;(2 )买麻藤目并不是被子植物的姊妹群而是松柏目的姊妹群 ,甚至网结于松柏类而成为松科的姊妹群。这些结果并不使人惊讶 ,因为对探讨像包含许多灭绝类群的种子植物系统 ,决不可能是仅仅单独应用现代类群资料所能完成的。假如生花植物支是成立的 ,但其名称以AGPB支代替生花植物支可能较为合理。     

被子植物花的起源:假说和证据

达尔文的 令人讨厌之谜 ,即被子植物的起源和早期演化 ,一直是植物系统学研究领域的热点 .被子植物区别于其它植物类群的一个显著特征就是花 ,因此 ,解决被子植物的起源之谜很大程度上取决于对被子植物花器官起源的研究 .对被子植物花器官的详尽研究已经在形态、解剖、古植物、形态发生、分子等方面积累了大量的证据 ,植物学家基于这些证据为被子植物花器官的起源提出了各种各样的解释 .综述了迄今为止被子植物花器官起源的主要学说流派 ,如 :真花学说、假花学说、生殖叶学说、生殖茎节学说、生花植物学说、新假花学说、古草本学说和 ANITA学说等 .根据研究手段和获得证据的方式 ,作者将被子植物花器官起源研究划分为 5个阶段 ,并简要阐述了各个阶段的代表学说和主要研究特点     

被子植物花的起源：假说和证据

达尔文的“令人讨厌之谜”,即被子植物的起源和早期演化,一直是植物系统学研究领域的热点。被子植物区别于其它植物类群的一个显著特征就是花,因此,解决被子植物的起源之谜很大程度上取决于对被子植物花器官起源的研究,对被子植物花器官的详尽研究已经在形态、解剖、古植物、形态发生、分子等方面积累了大量的证据,植物学家基于这些证据为被子植物花器官的起源提出了各种各样的解释。综述了迄今为止被子植物花器官起源的主要学说流派,如：真花学说、假花学说、生殖叶学说、生殖茎节学说、生花植物学说、新假花学说、古草本学说和ANITA学说等。根据研究手段和获得证据的方式。作者将被子植物花器官起源研究划分为5个阶段,并简要阐述了各个阶段的代表学说和主要研究特点。     

植物学通报2007年第24卷总目次

第1期(2007年1月)1植物进化发育生物学的形成与研究进展张剑,徐桂霞,薛皓月,胡瑾31花同源异型M A D S-b o x基因在被子植物中的功能保守性和多样性崔荣峰,孟征42花、基因、禾本科李贵生,陈明生49被子植物胚珠研究进展王自芬,任毅60MADS-box基因家族基因重复及其功能的多样性吕山花,孟征71形态性状、分子性状与同源性山红艳80黄连属(毛茛科)花的形态发生辜天琪,任毅87铁线莲属尾叶铁线莲组(毛茛科)基于形态学证据的分支系统学孙诚,谢磊,李良千99天然棕色棉纤维色素光谱学特性及其化学结构初步推断詹少华,林毅,蔡永萍,李正鹏105植物脂肪酸…     

生花植物概念简介及其名称的商榷

在上个世纪最后的20年里,系统学家应用形态性状对种子植物进行了大量的分支分析。其结果显示灭绝的五柱木属加上灭绝的本内苏铁目及尚存的买麻藤目是被子植物的姊妹群,形成一个强支,称之为生花植物支。生花植物假说对探讨被子植物起源有着重要影响,它激发人们讨论被子植物起源时间可能要提前到三叠纪甚至石炭纪,除了支持原有的真花学说外,还提出新恩格勒学说。但是,近年来对现存种子植物进行分子系统学研究的结果是：(1)拒绝接受生花植物概念;(2)买麻藤目并不是被子植物的姊妹群而是松柏目的姊妹群,甚至网结于松柏类而成为松科的姊妹群。这些结果并不使人惊讶,因为对探讨像包含许多灭绝类群的种子植物系统,决不可能是仅仅单独应用现代类群资料所能完成的。假如生花植物支是成立的,但其名称以AGPB支代替生花植物支可能较为合理。     

腺齿木科系统位置评述

腺齿木科(Trimeniaceae)含2属5种.形态学研究显示腺齿木科具有许多原始性状.最新的分子系统发育研究显示,腺齿木科是现存被子植物的重要基部类群之一.但有关腺齿木科的系统位置存在争议.被子植物(有花植物)的起源与辐射一直是植物系统学家关注的热点.对该科系统位置的研究历史与现状进行评述.     

The ABC model and its applicability to basal angiosperms

BACKGROUND: Although the flower is the central feature of the angiosperms, little is known of its origin and subsequent diversification. The ABC model has long been the unifying paradigm for floral developmental genetics, but it is based on phylogenetically derived eudicot models. Synergistic research involving phylogenetics, classical developmental studies, genomics and developmental genetics has afforded valuable new insights into floral evolution in general, and the early flower in particular. SCOPE AND CONCLUSIONS: Genomic studies indicate that basal angiosperms, and by inference the earliest angiosperms, had a rich tool kit of floral genes. Homologues of the ABCE floral organ identity genes are also present in basal angiosperm lineages; however, C-, E- and particularly B-function genes are more broadly expressed in basal lineages. There is no single model of floral organ identity that applies to all angiosperms; there are multiple models that apply depending on the phylogenetic position and floral structure of the group in question. The classic ABC (or ABCE) model may work well for most eudicots. However, modifications are needed for basal eudicots and, the focus of this paper, basal angiosperms. We offer 'fading borders' as a testable hypothesis for the basal-most angiosperms and, by inference, perhaps some of the earliest (now extinct) angiosperms.     

The multifaceted roles of FLOWERING LOCUS T in plant development

One of the key developmental processes in flowering plants is the differentiation of the shoot apical meristem into a floral meristem. This transition is regulated through the integration of environmental and endogenous stimuli, involving a complex, hierarchical signalling network. In arabidopsis, the FLOWERING LOCUS T (FT) protein, a mobile signal recognized as a major component of florigen, has a central position in mediating the onset of flowering. FT-like genes seem to be involved in regulating the floral transition in all angiosperms examined to date. Evidence from molecular evolution studies suggests that the emergence of FT-like genes coincided with the evolution of the flowering plants. Hence, the role of FT in floral promotion is conserved, but appears to be restricted to the angiosperms. Besides flowering, FT-like proteins have also been identified as major regulatory factors in a wide range of developmental processes including fruit set, vegetative growth, stomatal control and tuberization. These multifaceted roles of FT-like proteins have resulted from extensive gene duplication events, which occurred independently in nearly all modern angiosperm lineages, followed by sub- or neo-functionalization. This review assesses the plethora of roles that FT-like genes have acquired during evolution and their implications in plant diversity, adaptation and domestication.     

Reproduction and development in flowering plants]

Why mark the centenary of the independent discovery of double fertilization by Sergius Nawashin (1898) and Léon Guignard (1899), when biology has progressed so much since the beginning of the XXth century? This discovery still constitutes one of the key references in plant biology: double fertilization is unique to flowering plants among all living organisms. This meeting is also the occasion to associate angiosperm fertilization with developmental biology because of the localization of this event in the flower. Very important and significant progress has been made in elucidating flower development during the last ten years. And today it is possible to understand the diversity of floral structure present in the angiosperms in the context of a underlying mechanism of flower development inherited from their common ancestor. This special issue also allows a survey of these two broad scientific fields, plant reproduction and plant development (flower and embryo). It might also attract new, talented young scientists.     

The interplay between inflorescence development and function as the crucible of architectural diversity

Background

Most angiosperms present flowers in inflorescences, which play roles in reproduction, primarily related to pollination, beyond those served by individual flowers alone. An inflorescence''s overall reproductive contribution depends primarily on the three-dimensional arrangement of the floral canopy and its dynamics during its flowering period. These features depend in turn on characteristics of the underlying branching structure (scaffold) that supports and supplies water and nutrients to the floral canopy. This scaffold is produced by developmental algorithms that are genetically specified and hormonally mediated. Thus, the extensive inflorescence diversity evident among angiosperms evolves through changes in the developmental programmes that specify scaffold characteristics, which in turn modify canopy features that promote reproductive performance in a particular pollination and mating environment. Nevertheless, developmental and ecological aspects of inflorescences have typically been studied independently, limiting comprehensive understanding of the relations between inflorescence form, reproductive function, and evolution.

Scope

This review fosters an integrated perspective on inflorescences by summarizing aspects of their development and pollination function that enable and guide inflorescence evolution and diversification.

Conclusions

The architecture of flowering inflorescences comprises three related components: topology (branching patterns, flower number), geometry (phyllotaxis, internode and pedicel lengths, three-dimensional flower arrangement) and phenology (flower opening rate and longevity, dichogamy). Genetic and developmental evidence reveals that these components are largely subject to quantitative control. Consequently, inflorescence evolution proceeds along a multidimensional continuum. Nevertheless, some combinations of topology, geometry and phenology are represented more commonly than others, because they serve reproductive function particularly effectively. For wind-pollinated species, these combinations often represent compromise solutions to the conflicting physical influences on pollen removal, transport and deposition. For animal-pollinated species, dominant selective influences include the conflicting benefits of large displays for attracting pollinators and of small displays for limiting among-flower self-pollination. The variety of architectural components that comprise inflorescences enable diverse resolutions of these conflicts.     

The floral genome: an evolutionary history of gene duplication and shifting patterns of gene expression

Through multifaceted genome-scale research involving phylogenomics, targeted gene surveys, and gene expression analyses in diverse basal lineages of angiosperms, our studies provide insights into the most recent common ancestor of all extant flowering plants. MADS-box gene duplications have played an important role in the origin and diversification of angiosperms. Furthermore, early angiosperms possessed a diverse tool kit of floral genes and exhibited developmental 'flexibility', with broader patterns of expression of key floral organ identity genes than are found in eudicots. In particular, homologs of B-function MADS-box genes are more broadly expressed across the floral meristem in basal lineages. These results prompted formulation of the 'fading borders' model, which states that the gradual transitions in floral organ morphology observed in some basal angiosperms (e.g. Amborella) result from a gradient in the level of expression of floral organ identity genes across the developing floral meristem.     

中国昆虫生态学研究的透视

<正> 昆虫生态学在我国是通过几十年来密切结合农、林、牧实际,解决生产中不断提出的重大问题而发展起来的。随着研究问题不断深入,交叉学科不断渗透,使昆虫生态学在生物学宏观领域中成为迅速发展的一门学科。现将7个主要领域研究的动态加以评述。     

Floral phyllotaxis in basal angiosperms: development and evolution

To date, molecular developmental studies have focused on vegetative rather than floral phyllotaxis because vegetative shoot apices are technically more tractable than floral apices in model plants. In contrast to evolutionary changes in the phyllotaxis of vegetative shoots, however, changes in floral phyllotaxis appear to have played a major role in angiosperm evolution. Consolidation of a whorled floral phyllotaxis in derived groups allowed synorganization of floral organs and further adaptive radiations. In basal angiosperms, floral phyllotaxis is more flexible. To study these phenomena, we need clarification of the complex relations of both spiral and whorled phyllotaxis with divergence angles, plastochrons, spiral versus simultaneous initiation of organs, parastichies, orthostichies, organ series, and whorls. Improved resolution of phylogenetic relationships and increased knowledge of the diversity of floral phyllotaxis will allow us to trace evolutionary changes in floral phyllotaxis in ever more detail. Already, such surveys have confirmed that floral phyllotaxis was unusually labile early in angiosperm evolution. Whether the original floral phyllotaxis in angiosperms was spiral or whorled is equivocal, but it appears that spiral floral phyllotaxis in Magnoliales and Laurales is derived rather than primitive.     

Floral variation and floral genetics in basal angiosperms

Recent advances in phylogeny reconstruction and floral genetics set the stage for new investigations of the origin and diversification of the flower. We review the current state of angiosperm phylogeny, with an emphasis on basal lineages. With the surprising inclusion of Hydatellaceae with Nymphaeales, recent studies support the topology of Amborella sister to all other extant angiosperms, with Nymphaeales and then Austrobaileyales as subsequent sisters to all remaining angiosperms. Notable modifications from most recent analyses are the sister relationships of Chloranthaceae with the magnoliids and of Ceratophyllaceae with eudicots. We review "trends" in floral morphology and contrast historical, intuitive interpretations with explicit character-state reconstructions using molecular-based trees, focusing on (1) the size, number, and organization of floral organs; (2) the evolution of the perianth; (3) floral symmetry; and (4) floral synorganization. We provide summaries of those genes known to affect floral features that contribute to much of floral diversity. Although most floral genes have not been investigated outside of a few model systems, sufficient information is emerging to identify candidate genes for testing specific hypotheses in nonmodel plants. We conclude with a set of evo-devo case studies in which floral genetics have been linked to variation in floral morphology.