Perspective: the origin of flowering plants and their reproductive biology--a tale of two phylogenies

Recently, two areas of plant phylogeny have developed in ways that could not have been anticipated, even a few years ago. Among extant seed plants, new phylogenetic hypotheses suggest that Gnetales, a group of nonflowering seed plants widely hypothesized to be the closest extant relatives of angiosperms, may be less closely related to angiosperms than was believed. In addition, recent phylogenetic analyses of angiosperms have, for the first time, clearly identified the earliest lineages of flowering plants: Amborella, Nymphaeales, and a clade that includes Illiciales/ Trimeniaceae/Austrobaileyaceae. Together, the new seed plant and angiosperm phylogenetic hypotheses have major implications for interpretation of homology and character evolution associated with the origin and early history of flowering plants. As an example of the complex and often unpredictable interplay of phylogenetic and comparative biology, we analyze the evolution of double fertilization, a process that forms a diploid embryo and a triploid endosperm, the embryo-nourishing tissue unique to flowering plants. We demonstrate how the new phylogenetic hypotheses for seed plants and angiosperms can significantly alter previous interpretations of evolutionary homology and firmly entrenched assumptions about what is synapomorphic of flowering plants. In the case of endosperm, a solution to the century-old question of its potential homology with an embryo or a female gametophyte (the haploid egg-producing generation within the life cycle of a seed plant) remains complex and elusive. Too little is known of the comparative reproductive biology of extant nonflowering seed plants (Gnetales, conifers, cycads, and Ginkgo) to analyze definitively the potential homology of endosperm with antecedent structures. Remarkably, the new angiosperm phylogenies reveal that a second fertilization event to yield a biparental endosperm, long assumed to be an important synapomorphy of flowering plants, cannot be conclusively resolved as ancestral for flowering plants. Although substantive progress has been made in the analysis of phylogenetic relationships of seed plants and angiosperms, these efforts have not been matched by comparable levels of activity in comparative biology. The consequence of inadequate comparative biological information in an age of phylogenetic biology is a severe limitation on the potential to reconstruct key evolutionary historical events.     

Growth and development of conifer pollen tubes

Conifer pollen tubes are an important but underused experimental system in plant biology. They represent a major evolutionary step in male gametophyte development as an intermediate form between the haustorial pollen tubes of cycads and Ginkgo and the structurally reduced and faster growing pollen tubes of flowering plants. Conifer pollen grains are available in large quantities, most can be stored for several years, and they grow very well in culture. The study of pollen tube growth and development furthers our understanding of conifer reproduction and contributes towards our ability to improve on their productivity. This review covers taxonomy and morphology to cell, developmental, and molecular biology. It explores recent advances in research on conifer pollen and pollen tubes in vivo, focusing on pollen wall structure, male gametophyte development within the pollen wall, pollination mechanisms, pollen tube growth and development, and programmed cell death. It also explores recent research in vitro, including the cellular mechanisms underlying pollen tube elongation, in vitro fertilization, genetic transformation and gene expression, and pine pollen tube proteomics. With the ongoing sequencing of the Pinus taeda genome in several labs, we expect the use of conifer pollen tubes as an experimental system to increase in the next decade.     

植物开花逆转研究进展

介绍了植物开花逆转的概念,类型,发生条件,综述了在形态发生,生理,分子生物学等方面的研究进展,探讨了将开花逆转应用于植物发育生物学研究的可能性,作者提出,采用“诱导－非诱导－逆转”三位一体的实验系统,可提高植物光周期反应研究试验结果的可靠性,特别是可以提高在研究开花机理时的基因表达时空性方面的可靠性。     

DNA methylation dynamics during germline development

DNA methylation plays essential homeostatic functions in eukaryotic genomes. In animals, DNA methylation is also developmentally regulated and, in turn, regulates development. In the past two decades, huge research effort has endorsed the understanding that DNA methylation plays a similar role in plant development, especially during sexual reproduction. The power of whole-genome sequencing and cell isolation techniques, as well as bioinformatics tools, have enabled recent studies to reveal dynamic changes in DNA methylation during germline development. Furthermore, the combination of these technological advances with genetics, developmental biology and cell biology tools has revealed functional methylation reprogramming events that control gene and transposon activities in flowering plant germlines. In this review, we discuss the major advances in our knowledge of DNA methylation dynamics during male and female germline development in flowering plants.     

植物开花逆转研究进展

介绍了植物开花逆转的概念、类型、发生条件,综述了在形态发生、生理、分子生物学等方面的研究进展,探讨了将开花逆转应用于植物发育生物学研究的可能性。作者提出,采用“诱导—非诱导—逆转”三位一体的实验系统,可提高植物光周期反应研究试验结果的可靠性,特别是可以提高在研究开花机理时的基因表达时空性方面的可靠性。     

高等植物开花时程的基因调控(Ⅱ)

主要探讨如下几个问题:转基因植物中的拟南芥开花时程基因;拟南芥中的甲基化与实验胚胎学研究;高等植物开花时程控制的可能机制.目前本领域已成为植物发育分子生物学的前沿热点研究领域之一.结合有关工作,对这一世界性热门领域进行了系统的评述,希望能为国内同行提供有关参考,赶超世界的植物分子生物学先进水平,对分子水平与生物技术角度改良黄土高原生态环境有指导意义.     

Patterns of diversity of flower-visitor assemblages to the understory Araceae in a tropical mountain forest in Colombia

In the Araceae, pollination biology has been extensively evaluated in several genera mainly in lowland tropical areas. However, the influence of physical factors such as elevation or precipitation on plant/flower-visitor interactions at the community level remains challenging for this plant family, specifically in tropical mountain forests. The aim of this study was to analyze how the species diversity and abundance of flowering Araceae and their flower-visitor assemblages change along an elevation gradient and through time, in response to changes in mean monthly precipitation in a tropical mountain forest located in Jardín, Antioquia, Colombia. Eighteen flowering Araceae species and floral visitors belonging to seven orders were recorded. Flower-visitor diversity decreased with elevation. A trend of increasing flowering Araceae and flower-visitor abundance richness at intermediate levels of precipitation was observed. Diptera were dominant in the lower part and Thysanoptera were dominant in the upper part of the gradient. The pattern of plant species distribution along the elevation and the amount of precipitation influenced the availability of flower resources, and, as a consequence, affected the spatial and temporal composition of flower-visitor assemblages. Local strategies for the conservation of the diversity of insect flower-visitors and their interactions should be focused on the implementation of agricultural practices that reduce the use of pesticides within adjacent commercial plantations and the avoidance of illegal clearings, maintaining unbroken elevational gradients of forest, which is the only way to protect the flowering resources for anthophilous insects. At the same time, continuous forest promotes the maintenance of macro and microclimatic conditions, preserving the stability of insect populations and diversity amongst several functional groups.     

New dimensions for VIGS in plant functional genomics

Virus-induced gene silencing (VIGS) is an efficient tool for gene function studies. It has been used to perform both forward and reverse genetics to identify plant genes involved in several plant processes. However, this technology has not yet been used to its full potential. This can be attributed to several of its limitations such as inability to silence genes during seed germination and the non-stable nature of silencing. However, several recent studies have shown that these limitations can now be overcome. In this review, we will discuss the limitations of VIGS and suitable solutions. In addition, we also describe the recent improvements and future prospects of using VIGS in plant biology.     

A general stochastic model for sporophytic self-incompatibility

Disentangling the processes leading populations to extinction is a major topic in ecology and conservation biology. The difficulty to find a mate in many species is one of these processes. Here, we investigate the impact of self-incompatibility in flowering plants, where several inter-compatible classes of individuals exist but individuals of the same class cannot mate. We model pollen limitation through different relationships between mate availability and fertilization success. After deriving a general stochastic model, we focus on the simple case of distylous plant species where only two classes of individuals exist. We first study the dynamics of such a species in a large population limit and then, we look for an approximation of the extinction probability in small populations. This leads us to consider inhomogeneous random walks on the positive quadrant. We compare the dynamics of distylous species to self-fertile species with and without inbreeding depression, to obtain the conditions under which self-incompatible species can be less sensitive to extinction while they can suffer more pollen limitation.     

荒漠孑遗植物裸果木的开花物候特征

2014年和2015年连续2年对哈密和乌恰裸果木(Gymnocarpos przewalskii)自然种群的开花物候特征进行野外定点观测,并运用相对开花振幅、开花强度和开花同步性等参数对其开花物候特征进行分析。结果表明,2个种群每年均开1次花,4月下旬到5月上旬进入始花期,开花进程基本相似,均呈单峰曲线,具有较高的开花同步性,呈现"集中大量"开花模式;在个体水平上,始花期、终花期在种群和年份间略有差异;单花持续时间在年际间是稳定的,但在种群间有所不同,即乌恰种群的单花持续时间略长于哈密种群;不同裸果木种群的开花数与结籽数间均呈极显著正相关,开花同步性指数与相对开花强度间呈显著负相关;乌恰种群的始花期与开花数间呈负相关。裸果木个体开花振幅、开花同步指数及开花模式在种群和年份间的相似性很大程度上是由其在长期的地理变迁中形成的遗传特性决定的,而始花期和开花持续时间等开花物候特征的差异主要是由环境和气象因子(海拔、气温、降水及光照)差异造成。研究结果对于探讨该属植物的繁殖生物学特性及其保护具有重要意义。     

Fire-Stimulated Flowering: A Review and Look to the Future

The literature on fire-stimulated flowering is reviewed in terms of post-fire flowering patterns, proximal ultimate factors that affect flowering of plants post-fire, as well as application of evolutionary theory and development of evolutionary models in relation to fire-stimulated flowering. Consideration is given to the concepts involved, available empirical information, and areas that warrant further research. Understanding the factors responsible for fire-stimulated flowering is limited. Studies suggest factors involved, but confounding of variables prevents their separation. Experiments evaluate different factors, but there have been few such studies and little consideration regarding soil attributes, synergistic effects of multiple factors, and large-scale factors such as herbivory, florivory, seed dispersal, and pollination. Understanding the adaptive nature, and hence evolution, of fire-stimulated flowering is poor. Prevalence of particular fire-related responses has been related to geographical and other patterns, with deduction of adaptive nature of plant responses to fire and factors driving evolution of these responses. However, confounding of factors hinders this approach and can lead to disagreement between researchers. A better approach would be to hypothesize that evolution has resulted in plant responses to fire that are Evolutionarily Stable Strategies and to develop consequent mathematical models for such evolution, leading to predictions and tests. However, this approach requires quantitative assessment of responses to fire-related factors and influences of such responses on plant fitness, and so far few such assessments exist. This poor state of knowledge regarding plants with fire-stimulated flowering means that we cannot easily replace or enhance fire for managing species with fire-stimulated flowering, as sometimes necessary, and it also restricts their commercial cultivation. Experimental evaluation of plant responses to fire and development of evolutionary models is required. Our knowledge of the factors that trigger post-fire flowering by re-sprouting plant species remains poor, and so we cannot incorporate relationships between them and plant fitness into mathematical models of plant evolution.     

复粒稻种质资源及其遗传育种利用

水稻(Oryza sativa)是世界上最重要的粮食作物之一, 也是单子叶植物发育分子生物学研究的理想模式植物。穗部形态是影响水稻产量的重要因素, 也是当前水稻遗传改良和发育生物学研究的热点之一。复粒稻是发生于水稻穗粒部的一种突变体材料, 在形态上可分为小穗簇生型和颖壳多雌型两种, 有自然突变、杂交变异和理化诱变等多种来源途径。作为一种特异的水稻种质资源, 复粒稻在水稻新材料创建、复粒型新品种选育、改善杂交制种结实性及探讨单子叶植物成花机理等方面具有重要的研究价值, 对创造复粒新型育种材料、提高穗着粒密度及穗粒数、有效缩短穗长度、改良水稻植株性状及提高产量等都具有重要意义。该文综述了复粒稻的资源类型、来源途径、遗传分析、基因定位及其育种利用价值等方面的研究进展, 并提出了今后的研究方向, 以期为水稻遗传育种提供参考。     

Research on orchid biology and biotechnology

Orchidaceae constitute one of the largest families of angiosperms. They are one of the most ecological and evolutionary significant plants and have successfully colonized almost every habitat on earth. Because of the significance of plant biology, market needs and the current level of breeding technologies, basic research into orchid biology and the application of biotechnology in the orchid industry are continually endearing scientists to orchids in Taiwan. In this introductory review, we give an overview of the research activities in orchid biology and biotechnology, including the status of genomics, transformation technology, flowering regulation, molecular regulatory mechanisms of floral development, scent production and color presentation. This information will provide a broad scope for study of orchid biology and serve as a starting point for uncovering the mysteries of orchid evolution.     

Developmental phenotypic plasticity: Where ecology and evolution meet molecular biology

The plastic response of phenotypic traits to environmental change is a common research focus in several disciplines - from ecology and evolutionary biology to physiology and molecular genetics. The use of model systems such as the flowering plant Arabidopsis thaliana has facilitated a dialogue between developmental biologists asking how plasticity is controlled (proximate causes) and organismal biologists asking why plasticity exists (ultimate causes). Researchers studying ultimate causes and consequences are increasingly compelled to reject simplistic, ‘black box’ models, while those studying proximate causes and mechanisms are increasingly obliged to subject their interpretations to ecological ‘reality checks.’ We review the successful multidisciplinary efforts to understand the phytochrome-mediated shade-avoidance and light-seeking responses of flowering plants as a pertinent example of convergence between evolutionary and molecular biology. In this example, the two-way exchange between reductionist and holist camps has been essential to rapid and sustained progress. This should serve as a model for future collaborative efforts towards understanding the responses of organisms to their constantly changing environments.     

Plant chromatin: development and gene control

It is increasingly clear that chromatin is not just a device for packing DNA within the nucleus but also a dynamic material that changes as cellular environments alter. The precise control of chromatin modification in response to developmental and environmental cues determines the correct spatial and temporal expression of genes. Here, we review exciting discoveries that reveal chromatin participation in many facets of plant development. These include: chromatin modification from embryonic and meristematic development to flowering and seed formation, the involvement of DNA methylation and chromatin in controlling invasive DNA and in maintenance of epigenetic states, and the function of chromatin modifying and remodeling complexes such as SWI/SNF and histone acetylases and deacetylases in gene control. Given the role chromatin structure plays in every facet of plant development, chromatin research will undoubtedly be integral in both basic and applied plant biology.     

Fibonacci,quasicrystals and the beauty of flowers

The appearance of Fibonacci sequences and the golden ratio in plant structures is one of the great outstanding puzzles of biology. Here I suggest that quasicrystals, which naturally pack in the golden ratio, may be ubiquitous in biological systems and introduce the golden ratio into plant phyllotaxy. The appearance of golden ratio-based structures as beautiful indicates that the golden ratio may play a role in the development of consciousness and lead to the aesthetic natural selection of flowering plants.     

Challenges in studies on flowering time: interfaces between phenological research and the molecular network of flowering genes

Flowering time is a well-studied subject in ecology, evolution and molecular biology. Long-term phenological studies have shown relationships between flowering time and environmental and endogenous factors in many species. In contrast, molecular studies using model plants have revealed a complex regulatory network of flowering. We propose that flowering would be a model trait for the integrated study of ecology, evolution and molecular biology. We introduce briefly the flowering regulatory pathways of Arabidopsis thaliana, followed by molecular techniques such as transgenic manipulation, quantitative real-time PCR and detection of differentially expressed genes that could facilitate the study of ‘nonmodel’ species of ecological interest but with little available genome information. Application of the flowering gene network to wild species will be illustrated by two examples: modeling and prediction of the expression of flowering genes in Arabidopsis halleri, and the latitudinal cline of bud set and cessation in Populus. Finally, we discuss the challenges in integrating knowledge of the regulatory network on flowering into ecologically unique flowering phenomena such as synchronous intermittent mass flowering—the topic of this special issue.     

LTP在植物抗环境胁迫中的作用

脂质转移蛋(白Lipid transfer protein,LTP)是一类分泌蛋白,曾被认为是一种在体外膜间进行脂质转移的蛋白,因其作用对象很广泛,所以目前又称其为非特异性LTP(non-specific lipid transfer proteins,nsLTPs)。有很多证据表明nsLTPs可能参与多方面的植物细胞与生理生化反应,这些生物学功能包括:参与角质层的合成和胚胎的发育;适应各种胁迫环境;抗病原微生物等作用,尤其是在适应胁迫环境中,起到很大作用。介绍了nsLTPs在植物抗环境胁迫中所发挥的作用,同时对应用研究作了简单展望。     

The effects of fire and fragmentation on occurrence and flowering of a rare perennial plant

The pine rocklands of southern Florida are a fire-dependent forest associated with outcroppings of limestone. Pine rockland plants have several adaptations to fire, and for many species burns increase plant growth, flowering, and seedling establishment. The pine rockland forest has been reduced and fragmented in recent decades. Outside of Everglades National Park, only two percent of the original pine rocklands remain, and are in the form of small fragments. Habitat fragmentation may have a negative effect on the biology of plants; we investigated the effects of both fire and fragmentation on Angadenia berteroi (A.DC.) Miers, a threatened species of the southern Florida pine rockland. We estimated the density and flowering of A. berteroi using adaptive cluster sampling in six study sites with different fire and disturbance histories. A. berteroi is more abundant in the largest fragments, and those having experienced fire most recently. However, fragmentation and lack of fire did not appear to have a great impact on flowering or fruit production. Insights from this threatened species may provide impetus not only to conserve, but to properly manage remaining pine rocklands in south Florida.