The geographical relationships of the British and Irish flora: a comparison of pteridophytes, flowering plants, liverworts and mosses

Aim To compare the wider distribution of the pteridophytes, flowering plants, liverworts and mosses recorded as natives of Britain and Ireland, and to assess the proportion and distribution of species in each group which belong to predominantly extra-European genera. Location The species native to the British Isles are considered in relation to their distribution in the Northern Hemisphere, and predominantly extra-European genera are identified by consideration of the distribution and centre of numerical diversity of the genera worldwide. Methods A recent classification of the pteridophytes, flowering plants, liverworts and mosses into floristic elements is used in a quantitative study of the distribution of species in the four groups, the patterns of occurrence being compared by standardized residuals. Results Mosses and liverworts have more northerly distributions than the flowering plants in the Northern Hemisphere; pteridophytes show an intermediate pattern. There are marked differences in the longitudinal ranges of the groups south of the Boreal biome, with vascular plants tending to be confined to Europe or western Eurasia whereas cryptogams are more likely to occur additionally in North America or have a circumpolar distribution. The proportion of predominantly extra-European genera is particularly high in the pteridophytes and the liverworts and in these groups the rich hyperoceanic flora of the British Isles is almost entirely composed of species in genera that attain their centre of diversity in the tropics. Main conclusions Major taxonomic groups show different distribution patterns, reflecting their rates of evolution and powers of dispersal. The flowering plants, as the least mobile and most rapidly evolving group, have a greater proportion of their species in the European and Eurosiberian elements and a higher proportion of species from extra-European genera in the more northerly elements.     

The modern concepts on the primary cell walls of angiosperm plants

The modern view on polysaccharide composition of the primary cell wall of angiosperm plants was considered, and five classes of structural proteins were characterized. The model which shows location of enzyme complex synthesizing cellulose, as well as structural model of primary cell walls of the most flowering plants were presented. Besides, there were shown cell wall reactions to environmental factors which have contradictory character and have to be deeply and thoroughly analyzed.     

高等植物开花时程的基因调控（Ⅰ）

高等植物从营养生长向生殖生长及发育转变的时程具有重要意义,但是了解得很少。近6年来利用分子遗传学方法详细地分析了拟南芥中的这一转变的时程变化,为高等植物开花时程的基因调控提供了一个很好的模式。有关早期或晚期开花表现型的大量突变体及遗传变异得到了阐述。这里谈到的表现型对影响开花转变的环境及内部因子的控制有重大作用。通过分子生物学、遗传学和生理学分析已经鉴定了参与此过程的不同组分,如光识别和昼夜节律（circadian rhythm）因子。另外,通过克隆某些花诱导基因及其相应的靶基因已经对参与开花信号转导途径（signal transduction pathway）的相关因子进行了系统的鉴定,这些开创性工作大大促进了高等植物开花时程的基因表达调控研究及其机理的阐明。本实验室在以黄瓜、新红宝西瓜、西葫芦为材料所获得的部分结果基础上,主要以近六年来在拟南芥方面获得的进展为依据,对高等植物开花时程的基因调控作一系统的总结,并对其开花时程基因调控的机理提出可能的作用理论模型。     

Life history variation and infraspecific heterochrony inRhinanthus glacialis (Scrophulariaceae)

Comparative cultivation experiments were made with 14 populations of the annual, hemiparasitic speciesRhinanthus glacialis from seven different habitat types. Phenological data were recorded and plants from field samples and from corresponding experimental samples were morphologically analysed.There are up to ten weeks difference in the start of flowering between means of the extreme experimental populations. Flowering time variation correlates with the number of internodes and also with most of the other morphological characters. Flowering time also shows close relationships to different grassland types and the examined populations appear to be well-adapted to the environmental conditions in natural or man-influenced grassland types as far as flowering time is concerned. The number of internodes is mostly insensitive to the applied experimental influences. The rate of internode production is constant. Trade-offs are suggested for some morphological traits (i.e., few internodes with few branches and thus few flowers as trade-off for early flowering). — Previous studies of life history variation based on population data have not been incorporated into the theory of heterochrony. The pattern of life history variation of populations ofR. glacialis together with information about morphology, ecology, and phylogeny suggests that some variants have evolved from ancestors via heterochronic processes. Earlier flowering populations from montane hay meadows, litter meadows, and dry meadows belong to paedomorphic variants. These have fewer internodes and branches as well as reduced height in comparison with the suggested ancestor. Later flowering populations from subalpine limestone grassland belong to a peramorphic variant with more internodes, more branches, and taller stems in comparison with the suggested ancestor.     

引种保护植物对西双版纳极端冷年的春季物候响应

气温被普遍认为是春季物候期最主要的控制因子之一,然而低温对植物物候的影响效应一直都存在不同的观点。西双版纳由于地处热带地区的北缘,其气温相对于赤道附近的热带地区较低。自1959年以来,西双版纳热带植物园引入了来自世界各个热带地区的4万余种植物进行保护,之前的研究证明西双版纳的低温对这些引种植物的生长有很大影响。因此,1974年西双版纳出现的极端低温势必对引种植物造成极大威胁,同时也是对这些植物低温适应能力的一个考验。通过对比43种引种植物物候期（生长抽梢期与开花期）在1974年与常年的差异情况,分析不同来源（热带亚洲、热带美洲与热带非洲）引种植物对西双版纳低温的适应性。结果表明,经历西双版纳1974年初的极端低温之后,使81%的引种植物生长抽梢期提前,同时也造成35%的引种植物在该年没有开花;而植物生长抽梢提前的主要原因则是极端低温以及低温过后气温迅速回升。引种植物均能顺利度过1974年的最冷时期,并出现生长抽梢物候,这意味着引种植物在经历极端低温之后都能够进行正常的生长活动,但极端低温对引种植物繁殖活动的不利影响大于其对生长活动的影响;引种植物对西双版纳极端低温的适应能力由大到小顺序依次为：亚洲来源植物〉美洲来源植物〉非洲来源植物。因此在迁地保护植物的选择过程中,应多选择亚洲热带植物,其次为美洲热带植物,而对非洲热带植物的引入则需谨慎考察。     

Young but not relatively old retrotransposons are preferentially located in gene‐rich euchromatic regions in tomato (Solanum lycopersicum) plants

Long terminal repeat (LTR) retrotransposons are the major DNA components of flowering plants. They are generally enriched in pericentromeric heterochromatin regions of their host genomes, which could result from the preferential insertion of LTR retrotransposons and the low effectiveness of purifying selection in these regions. To estimate the relative importance of the actions of these two factors on their distribution pattern, the LTR retrotransposons in Solanum lycopersicum (tomato) plants were characterized at the genome level, and then the distribution of young elements was compared with that of relatively old elements. The current data show that old elements are mainly located in recombination‐suppressed heterochromatin regions, and that young elements are preferentially located in the gene‐rich euchromatic regions. Further analysis showed a negative correlation between the insertion time of LTR retrotransposons and the recombination rate. The data also showed there to be more solo LTRs in genic regions than in intergenic regions or in regions close to genes. These observations indicate that, unlike in many other plant genomes, the current LTR retrotransposons in tomatoes have a tendency to be preferentially located into euchromatic regions, probably caused by their severe suppression of activities in heterochromatic regions. These elements are apt to be maintained in heterochromatin regions, probably as a consequence of the pericentromeric effect in tomatoes. These results also indicate that local recombination rates and intensities of purifying selection in different genomic regions are largely responsible for structural variation and non‐random distribution of LTR retrotransposons in tomato plants.     

Problem of the structural and functional organization of the immature motor center

A hypothetical structural and functional scheme of organization of the immature spinal motor center is proposed, based on our own studies carried out on rat pups of the first month of postnatal development as well as on the analysis of data from literature. Taking into account peculiarities of functioning of various chains of the segmental reflex apparatus (high excitability of the motor center, heterochrony in the development of interneurons of the spinal cord dorsal horns as well as of excitatory and inhibitory mechanisms, possibility of generation of rhythmic activity by one of the half-centers of the motor generator regardless of the activity of the other one) and the mechanisms of its cholinergic and catecholaminergic regulation, age-related changes are considered, which are connected with the organization of interrelations of elements both within the motor center and from the descending regulatory systems.     

Regulation of flowering plants of different biotypes

The patterns of control of flowering are analyzed in plants of different biotypes. The photoperiodic reaction of flowering taken as an example, the whole net of control is considered: from the environmental stimulus through its physiological transformation in the leaf in the corresponding hormonal impulse which, in turn, controls the realization of genetic programme and formation of generative organs in the stem buds. The photoperiodically neutral plants taken as an example, the patterns of age control of flowering are considered. In plants of different photoperiodic groups the synthesis of complementary components of florigen was shown to proceed either autonomously under the photoperiodic effect or under the inducing effect of definite photoperiods. The autonomous and inducible mechanisms of biosynthesis of the flowering hormones have a common base, the genetic system to which the environment sends its stimuli through the hormonal interactions. The interaction of hormonal and genetic developmental factors is considered, the evocation of flowering in the stem buds taken as an example.     

引种保护植物对西双版纳极端冷年的春季物候响应

气温被普遍认为是春季物候期最主要的控制因子之一, 然而低温对植物物候的影响效应一直都存在不同的观点。西双版纳由于地处热带地区的北缘, 其气温相对于赤道附近的热带地区较低。自1959年以来, 西双版纳热带植物园引入了来自世界各个热带地区的4万余种植物进行保护, 之前的研究证明西双版纳的低温对这些引种植物的生长有很大影响。因此,1974年西双版纳出现的极端低温势必对引种植物造成极大威胁, 同时也是对这些植物低温适应能力的一个考验。通过对比43种引种植物物候期(生长抽梢期与开花期)在1974年与常年的差异情况, 分析不同来源(热带亚洲、热带美洲与热带非洲)引种植物对西双版纳低温的适应性。结果表明, 经历西双版纳1974年初的极端低温之后, 使81%的引种植物生长抽梢期提前, 同时也造成35%的引种植物在该年没有开花; 而植物生长抽梢提前的主要原因则是极端低温以及低温过后气温迅速回升。引种植物均能顺利度过1974年的最冷时期, 并出现生长抽梢物候, 这意味着引种植物在经历极端低温之后都能够进行正常的生长活动, 但极端低温对引种植物繁殖活动的不利影响大于其对生长活动的影响; 引种植物对西双版纳极端低温的适应能力由大到小顺序依次为: 亚洲来源植物>美洲来源植物>非洲来源植物。因此在迁地保护植物的选择过程中, 应多选择亚洲热带植物, 其次为美洲热带植物, 而对非洲热带植物的引入则需谨慎考察。     

Nonheterochronic developmental changes underlie morphological heterochrony in the evolution of the Ardeidae

Evolutionary changes in developmental timing and rates (heterochrony) are a source of morphological variation. Here we explore a central issue in heterochronic analysis: are the alterations in developmental timing and rates the only factor underlying morphological heterochrony? Tarsometatarsal growth through endochondral ossification in Ardeidae evolution has been taken as a case study. Evolutionary changes in bone growth rate (morphological heterochrony) might be either (a) the result of alterations in the mitotic frequency of epiphyseal chondrocytes (process‐heterochrony hypothesis), or (b) the outcome of alterations in the number of proliferating cells or in the size of hypertrophic chondrocytes (structural hypothesis). No correlation was found between tarsometatarsal growth rates and the frequency of cell division. However, bone growth rates were significantly correlated with the number of proliferating cells. These results support the structural hypothesis: morphological acceleration and deceleration are the outcome of evolutionary changes in one structural variable, the number of proliferating cells.     

Stochastic flowering phenology in Dactylis glomerata populations described by Markov chain modelling

Understanding the relationship between flowering patterns and pollen dispersal is important in climate change modelling, pollen forecasting, forestry and agriculture. Enhanced understanding of this connection can be gained through detailed spatial and temporal flowering observations on a population level, combined with modelling simulating the dynamics. Species with large distribution ranges, long flowering seasons, high pollen production and naturally large populations can be used to illustrate these dynamics. Revealing and simulating species-specific demographic and stochastic elements in the flowering process will likely be important in determining when pollen release is likely to happen in flowering plants. Spatial and temporal dynamics of eight populations of Dactylis glomerata were collected over the course of two years to determine high-resolution demographic elements. Stochastic elements were accounted for using Markov chain approaches in order to evaluate tiller-specific contribution to overall population dynamics. Tiller-specific developmental dynamics were evaluated using three different RV matrix correlation coefficients. We found that the demographic patterns in population development were the same for all populations with key phenological events differing only by a few days over the course of the seasons. Many tillers transitioned very quickly from non-flowering to full flowering, a process that can be replicated with Markov chain modelling. Our novel approach demonstrates the identification and quantification of stochastic elements in the flowering process of D. glomerata, an element likely to be found in many flowering plants. The stochastic modelling approach can be used to develop detailed pollen release models for Dactylis, other grass species and probably other flowering plants.

     

FTIP1 is an essential regulator required for florigen transport

The capacity to respond to day length, photoperiodism, is crucial for flowering plants to adapt to seasonal change. The photoperiodic control of flowering in plants is mediated by a long-distance mobile floral stimulus called florigen that moves from leaves to the shoot apex. Although the proteins encoded by FLOWERING LOCUS T (FT) in Arabidopsis and its orthologs in other plants are identified as the long-sought florigen, whether their transport is a simple diffusion process or under regulation remains elusive. Here we show that an endoplasmic reticulum (ER) membrane protein, FT-INTERACTING PROTEIN 1 (FTIP1), is an essential regulator required for FT protein transport in Arabidopsis. Loss of function of FTIP1 exhibits late flowering under long days, which is partly due to the compromised FT movement to the shoot apex. FTIP1 and FT share similar mRNA expression patterns and subcellular localization, and they interact specifically in phloem companion cells. FTIP1 is required for FT export from companion cells to sieve elements, thus affecting FT transport through the phloem to the SAM. Our results provide a mechanistic understanding of florigen transport, demonstrating that FT moves in a regulated manner and that FTIP1 mediates FT transport to induce flowering.     

被子植物叶绿体基因组的结构变异研究进展

叶绿体是绿色植物特有的细胞器,其基因组信息被广泛应用于植物系统发育和比较基因组学研究。目前,越来越多的物种有了叶绿体全基因组序列,人们对叶绿体基因组的结构及其变异规律有了更深入的了解。该文对近年来国内外有关被子植物叶绿体基因组插入/缺失、短片段倒位与重复、基因组结构重排以及基因丢失等结构变异式样的研究进展进行综述,并分析了叶绿体基因组结构研究中仍存在的问题以及该领域未来的发展趋势。     

MicroRNAs in the moss Physcomitrella patens

Having diverged from the lineage that lead to flowering plants shortly after plants have established on land, mosses, which share fundamental processes with flowering plants but underwent little morphological changes by comparison with the fossil records, can be considered as an evolutionary informative place. Hence, they are especially useful for the study of developmental evolution and adaption to life on land. The transition to land exposed early plants to harsh physical conditions that resulted in key physiological and developmental changes. MicroRNAs (miRNAs) are an important class of small RNAs (sRNAs) that act as master regulators of development and stress in flowering plants. In recent years several groups have been engaged in the cloning of sRNAs from the model moss Physcomitrella patens. These studies have revealed a wealth of miRNAs, including novel and conserved ones, creating a unique opportunity to broaden our understanding of miRNA functions in land plants and their contribution to the latter??s evolution. Here we review the current knowledge of moss miRNAs and suggest approaches for their functional analysis in P. patens.     

Patterns of floral construction in ontogeny and phylogeny

Among the flowering plants flowers are relatively closed systems of a considerable complexity and therefore more suitable for comparative developmental studies than other organs. They work with a few kinds of structural elements that occur in variable or fixed numbers and are arranged in patterns more or less constrained by developmental conditions. Diversity of floral structure occurs at different levels, with emphasis either on number and arrangement or on synorganization and form of the structural elements. The first level is shown here with examples of several representatives of the primitive subclasses Magnoliidae and Hamamelididae, the second with Asclepiadaceae representing the most advanced subclass of dicotyledons Asteridae. Although early ontogenetic patterns are fairly constant among larger groups, especially the sequence of initiation and position of the different classes of structural elements, there is no strict stability in any of the parameters studied.     

Evolution of unusual morphologies in Lentibulariaceae (bladderworts and allies) and Podostemaceae (river-weeds): a pictorial report at the interface of developmental biology and morphological diversification

Background Various groups of flowering plants reveal profound (‘saltational’) changes of their bauplans (architectural rules) as compared with related taxa. These plants are known as morphological misfits that appear as rather large morphological deviations from the norm. Some of them emerged as morphological key innovations (perhaps ‘hopeful monsters’) that gave rise to new evolutionary lines of organisms, based on (major) genetic changes.Scope This pictorial report places emphasis on released bauplans as typical for bladderworts (Utricularia, approx. 230 secies, Lentibulariaceae) and river-weeds (Podostemaceae, three subfamilies, approx. 54 genera, approx. 310 species). Bladderworts (Utricularia) are carnivorous, possessing sucking traps. They live as submerged aquatics (except for their flowers), as humid terrestrials or as epiphytes. Most Podostemaceae are restricted to rocks in tropical river-rapids and waterfalls. They survive as submerged haptophytes in these extreme habitats during the rainy season, emerging with their flowers afterwards. The recent scientific progress in developmental biology and evolutionary history of both Lentibulariaceae and Podostemaceae is summarized.Conclusions Lentibulariaceae and Podostemaceae follow structural rules that are different from but related to those of more typical flowering plants. The roots, stems and leaves – as still distinguishable in related flowering plants – are blurred (‘fuzzy’). However, both families have stable floral bauplans. The developmental switches to unusual vegetative morphologies facilitated rather than prevented the evolution of species diversity in both families. The lack of one-to-one correspondence between structural categories and gene expression may have arisen from the re-use of existing genetic resources in novel contexts. Understanding what developmental patterns are followed in Lentibulariaceae and Podostemaceae is a necessary prerequisite to discover the genetic alterations that led to the evolution of these atypical plants. Future molecular genetic work on morphological misfits such as bladderworts and river-weeds will provide insight into developmental and evolutionary aspects of more typical vascular plants.     

Selection of floral resources to optimise conservation of agriculturally-functional insect groups

Non-crop vegetation in agricultural landscapes can provide a means of conserving insects in farmed landscapes and optimising on-farm ecosystem services as a result. Inclusion of floral resources may be particularly useful in conserving many beneficial insects, where groups including pollinators and pest natural enemies often rely on nectar and pollen during (at least part of) their life-cycle. As not all flowers are equally suited to all beneficial insects, selection of appropriate flowering plants is key to ensuring that conservation targets are met and benefits to ecosystems services realised as a result. This short paper describes an experiment conducted to assess the ‘total temporal attractiveness’ of a range of British wildflowers to selected functional insect groups. The results obtained demonstrate that flowering period alone is a poor indicator of plant suitability to insects, where no relationship existed between this and attraction to insects overall. Data also suggest that, based on attraction over a season, certain flowering plants are more likely to be of general insect conservation value and/or benefit to functional insect groups than others. Attraction to pest insects was also considered, with relatively high catches of thrips and pollen beetles observed in flowering stands of some plants.     

Antifungal proteins and peptides of leguminous and non-leguminous origins

Antifungal proteins and peptides, as their names imply, serve a protective function against fungal invasion. They are produced by a multitude of organisms including leguminous flowering plants, non-leguminous flowering plants, gymnosperms, fungi, bacteria, insects and mammals. The intent of the present review is to focus on the structural and functional characteristics of leguminous, as well as non-leguminous, antifungal proteins and peptides. A spectacular diversity of amino acid sequences has been reported. Some of the antifungal proteins and peptides are classified, based on their structures and/or functions, into groups including chitinases, glucanases, thaumatin-like proteins, thionins, and cyclophilin-like proteins. Some of the well-known proteins such as lectins, ribosome inactivating proteins, ribonucleases, deoxyribonucleases, peroxidases, and protease inhibitors exhibit antifungal activity. Different antifungal proteins may demonstrate different fungal specificities. The mechanisms of antifungal action of only some antifungal proteins including thaumatin-like proteins and chitinases have been elucidated.     

影响开花植物物种多样化速率变化的因素探讨

开花植物(被子植物)是目前最为繁荣兴盛的陆地植物,是地球上种类最多、分布最广、适应性最强的优势植物类群。然而,物种多样性在开花植物的谱系中极端不均匀,同时其物种并非匀速增加,而是集中在某些时期快速扩张。这种现象暗示了开花植物的多样性受到某些因素的调控,而揭示影响开花植物多样化速率的机制便是这数十年来植物学家孜孜不倦努力的目标。该文介绍了当前的研究进展。研究显示,那些与类群分类相关的关键性状大多与多样化无关,而早期被视为决定因素的花性状,目前被认为影响较低或是需要多性状联合才产生实际影响。多倍化是一个受到高度关注的候选因素,被认为能提供物种扩张所需的遗传物质,在适当的情况或其他因素的参与下促成物种多样化。然而,许多研究也呈现各类的争议,这些悖论或许代表了多倍化不是决定因素,而是扮演着间接的角色。另外,环境的变异(温度、湿度、空间)也能提高多样性的发生。整体而言,影响开花植物多样化速率的原因,很可能是一种综合因素共同促成的结果,而每个植物类群多样化速率提升的原因亦可能不尽相同。因此,先分别对植物类群进行探讨,再进行整合,或许更能提供一个确实且普遍的模型。     

Environmental regulation of flowering

The timing of flower initiation is a highly plastic developmental process. To achieve reproductive success, plants must select the most favourable season to initiate reproductive development; this in turn requires continuous monitoring of environmental factors and a properly response. Environmental factors which change in a predictable fashion along the year, such as light and temperature, are the most relevant in terms of selection of the flowering season. In Arabidopsis and more recently in a few other species, molecular genetic analyses are providing a way to identify the genes involved in the regulation of flowering time. From gene sequences it is possible to develop hypotheses regarding molecular function and to infer some of the molecular mechanisms involved in the environmental regulation of flowering time. In this paper, we summarize recent discoveries concerning the mechanisms which plants use to perceive and respond to major environmental factors (light and temperature) and their different components. We focus mainly on annual plants and especially on Arabidopsis because most of the available molecular and functional data come from this species. However, additional information arising from other plant systems is also considered.