生长素对拟南芥叶片发育调控的研究进展

叶片（包括子叶）是茎端分生组织产生的第一类侧生器官,在植物发育中具有重要地位。早期叶片发育包括三个主要过程：叶原基的起始,叶片腹背性的建立和叶片的延展。大量证据表明叶片发育受到体内遗传机制和体外环境因子的双重调节。植物激素,尤其是生长素在协调体内外调节机制中起着不可或缺的作用。生长素的稳态调控、极性运输和信号转导影响叶片发育的全过程。本文着重介绍生长素在叶片生长发育和形态建成中的调控作用,试图了解复杂叶片发育调控网络。     

Cellular Differentiation and Leaf Morphogenesis in Arabdopsis

A focused approach that exploits a single plant species, namely, Arabidopsis thaliana, as a means to understand how leaf cells differentiate and the factors that govern overall leaf morphogenesis has begun to generate a significant body of knowledge in this model plant. Although many studies have concentrated on specific cell types and factors that control their differentiation, some degree of consensus is starting to be reached. However, an understanding of specific mechanisms by which cells differentiate in relation to their position, that appears to be an overriding factor in this process, is not yet in place for cell types in the Arabidopsis leaf. It is clear that perturbations in cellular development within the leaf do not necessarily have a general effect on morphogenesis. Environmental factors, particularly light, have been known to affect leaf cell differentiation and expansion, and endogenous hormones also appear to play an important role, through mechanisms that are beginning to be uncovered. It is likely that continued identification of genes involved in leaf development and their regulation in relation to positional information or other cues will lead to a clearer understanding of the control of differentiation and morphogenesis in the Arabidopsis leaf.     

Plant architecture

Plant architecture is species specific, indicating that it is under strict genetic control. Although it is also influenced by environmental conditions such as light, temperature, humidity and nutrient status, here we wish to focus only on the endogenous regulatory principles that control plant architecture. We summarise recent progress in the understanding of the basic patterning mechanisms involved in the regulation of leaf arrangement, the genetic regulation of meristem determinacy, i.e. the decision to stop or continue growth, and the control of branching during vegetative and generative development.  Finally, we discuss the basis of leaf architecture and the role of cell division and cell growth in morphogenesis.     

生长素对拟南芥叶片发育调控的研究进展

叶片(包括子叶)是茎端分生组织产生的第一类侧生器官, 在植物发育中具有重要地位。早期叶片发育包括三个主要过程: 叶原基的起始, 叶片腹背性的建立和叶片的延展。大量证据表明叶片发育受到体内遗传机制和体外环境因子的双重调节。植物激素, 尤其是生长素在协调体内外调节机制中起着不可或缺的作用。生长素的稳态调控、极性运输和信号转导影响叶片发育的全过程。本文着重介绍生长素在叶片生长发育和形态建成中的调控作用, 试图了解复杂叶片发育调控网络。     

玉米叶形相关性状的Meta-QTL及候选基因分析

叶长、叶宽、叶面积及叶夹角不仅影响玉米(Zea mays)光合效率, 也是株型的重要构成因素。通过对620个叶形QTL进行整合, 构建不同遗传背景下的叶形QTL整合图谱, 利用元分析发掘出22个叶长、22个叶宽、12个叶面积以及17个叶夹角mQTL; 进一步运用生物信息学手段, 确定44个与叶片发育密切相关的候选基因。分析发现, 仅有NAL7-like、YABBY6- like和GRF2等13个基因位于mQTL区间内, 而玉米中已克隆的KNOTTED1、AN3/GIF1、rgd1/lbl1、mwp1及SRL2-like、HYL1-like和CYCB2;4-like等水稻(Oryza sativa)和拟南芥(Arabidopsis thaliana)叶形同源基因位于未被整合的QTL内; 对44个候选基因在叶片长、宽、厚发育过程中基部-末端、中央-边缘、远轴-近轴的调控机理进行归纳分析, 发现玉米中仅有少数几个候选基因被报道, 揭示了叶形发育的部分分子机理。因此, 对玉米叶形相关mQTL/QTL及基因进行全面深入的分析, 不仅有助于增加对其遗传结构的了解, 发掘更多候选基因, 阐明叶形发育和形成的分子机制, 还可为耐密理想株型的分子标记辅助选择提供依据。     

Leaf canopy as a dynamic system: ecophysiology and optimality in leaf turnover

BACKGROUND AND AIMS: In a leaf canopy, there is a turnover of leaves; i.e. they are produced, senesce and fall. These processes determine the amount of leaf area in the canopy, which in turn determines canopy photosynthesis. The turnover rate of leaves is affected by environmental factors and is different among species. This mini-review discusses factors responsible for leaf dynamics in plant canopies, focusing on the role of nitrogen. SCOPE: Leaf production is supported by canopy photosynthesis that is determined by distribution of light and leaf nitrogen. Leaf nitrogen determines photosynthetic capacity. Nitrogen taken up from roots is allocated to new leaves. When leaves age or their light availability is lowered, part of the leaf nitrogen is resorbed. Resorbed nitrogen is re-utilized in new organs and the rest is lost with dead leaves. The sink-source balance is important in the regulation of leaf senescence. Several models have been proposed to predict response to environmental changes. A mathematical model that incorporated nitrogen use for photosynthesis explained well the variations in leaf lifespan within and between species. CONCLUSION: When leaf turnover is at a steady state, the ratio of biomass production to nitrogen uptake is equal to the ratio of litter fall to nitrogen loss, which is an inverse of the nitrogen concentration in dead leaves. Thus nitrogen concentration in dead leaves (nitrogen resorption proficiency) and nitrogen availability in the soil determine the rate of photosynthesis in the canopy. Dynamics of leaves are regulated so as to maximize carbon gain and resource-use efficiency of the plant.     

Shade avoidance and the regulation of leaf inclination in Arabidopsis

As a rosette plant, Arabidopsis thaliana forms leaves near to the ground, which causes the plant to be vulnerable to shading by neighbours. One mechanism to avoid such shading is the regulation of leaf inclination, such that leaves can be raised to more vertical orientations to prevent neighbouring leaves from overtopping them. Throughout Arabidopsis rosette development, rosette leaves move to more vertical orientations when shaded by neighbouring leaves, exposed to low light levels or placed in the dark. After dark-induced reorientation of leaves, returning them to white light causes the leaves to reorient to more horizontal inclinations. These light-dependent leaf movements are more robust than, and distinct from, the diurnal movements of rosette leaves. However, the movements are gated by the circadian clock. The light-dependent leaf orientation response is mediated primarily through phytochromes A, B and E, with the orientation varying with the ratio of red light to far-red light, consistent with other shade-avoidance responses. However, even plants lacking these phytochromes were able to alter leaf inclination in response to white light, suggesting a role for other photoreceptors. In particular, we found significant changes in leaf inclination for plants exposed to green light. This green light response may be caused, in part, by light-dependent regulation of abscisic acid (ABA) biosynthesis.     

植物叶片最大羧化速率及其对环境因子响应的研究进展

植物叶片最大羧化速率对环境因子的响应关系是陆地生态系统生产力与碳收支研究的重要方面。论文从测定方法、影响因子与模拟模型3方面综述了植物叶片最大羧化速率及其对环境因子响应研究的最新进展,指出现有的植物叶片最大羧化速率对单个环境因子的响应研究严重制约着陆地生态系统生产力的准确评估。为弄清植物叶片最大羧化速率对环境因子的综合响应关系,迫切需要加强以下研究:(1)植物叶片最大羧化速率的生物与环境控制机制研究;(2)生物与环境因子协同作用下的植物叶片最大羧化速率定量模拟及其尺度化研究;(3)植物叶片最大羧化速率的环境因子阈值研究。     

The rice <Emphasis Type="Italic">bright green leaf</Emphasis> (<Emphasis Type="Italic">bgl</Emphasis>) locus encodes OsRopGEF10, which activates the development of small cuticular papillae on leaf surfaces

Development of specialized epidermal cells and structures plays a key role in plant tolerance to biotic and abiotic stresses. In the paddy field, the bright green leaf (bgl) mutants of rice (Oryza sativa) exhibit a luminous green color that is clearly distinguishable from the normal green of wild-type plants. Transmission and scanning electron microscopy revealed that small cuticular papillae (or small papillae; SP), nipple-like structures, are absent on the adaxial and abaxial leaf surfaces of bgl mutants, leading to more direct reflection and less diffusion of green light. Map-based cloning revealed that the bgl locus encodes OsRopGEF10, one of eleven OsRopGEFs in rice. RopGEFs (guanine nucleotide exchange factors for Rop) activate Rop/Rac GTPases, acting as molecular switches in eukaryotic signal transduction by replacing the bound GDP (inactive form) with GTP (active form) in response to external or internal cues. In agreement with the timing of SP initiation on the leaf epidermis, OsRopGEF10 is most strongly expressed in newly developing leaves before emergence from the leaf sheath. In yeast two-hybrid assays, OsRopGEF10 interacts with OsRac1, one of seven OsRac proteins; consistent with this, both proteins are localized in the plasma membrane. These results suggest that OsRopGEF10 activates OsRac1 to turn on the molecular signaling pathway for SP development. Together, our findings provide new insights into the molecular genetic mechanism of SP formation during early leaf morphogenesis.     

植物叶缘形态的发育调控机理

生物多样性研究的关键问题之一是表型多样性的形成和演化机制, 因为表型多样性与物种多样性密切相关, 同时又承载着遗传和环境的变异信息。植物的叶具有丰富的形态多样性, 而叶形多样性很大程度上体现在叶边缘形态的变异。叶边缘的形态可从全缘、锯齿状到具有不同程度(深浅)和不同式样(羽状或掌状、回数等)的裂片(在发育研究中复叶的小叶也描述为裂片)。关于叶缘齿/裂的发育调控机制, 在拟南芥(Arabidopsis thaliana)、碎米荠(Cardamine hirsuta)、番茄(Solanum lycopersicum)等模式植物中已有较深入的探讨。研究发现, 很多转录因子、小分子RNA及植物激素对叶齿/裂或小叶的形成具有调控作用, 其中生长素输出途径中的转录因子NAM/CUC、miR164以及高浓度生长素的反馈调控可能起到核心作用, 而且该调控模块在真双子叶植物中较为保守; TCP类、SPL类转录因子和其他一些miRNA也在生长素输出途径中发挥作用; 关于KNOX家族转录因子的作用, 虽然多数研究是围绕复叶的形态建成, 但也有数据显示其在叶裂发育中发挥作用。此外, 对拟南芥和碎米荠等十字花科植物的研究还发现, 调控基因RCO通过抑制小叶/裂片之间的细胞增殖而对小叶/叶裂的发育发挥作用。本文综述上述多角度的研究进展, 并尝试概括叶边缘形态的发育调控网络, 为关于叶缘形态多样性形成机制的研究提供可参考的切入点。     

ROP GTPase-mediated auxin signaling regulates pavement cell interdigitation in Arabidopsis thaliana

In multicellular plant organs, cell shape formation depends on molecular switches to transduce developmental or environmental signals and to coordinate cell‐to‐cell communication. Plants have a specific subfamily of the Rho GT Pase family, usually called Rho of Plants(ROP), which serve as a critical signal transducer involved in many cellular processes. In the last decade, important advances in the ROP‐mediated regulation of plant cell morphogenesis have been made by using Arabidopsis thaliana leaf and cotyledon pavement cells.Especially, the auxin‐ROP signaling networks have been demonstrated to control interdigitated growth of pavement cells to form jigsaw‐puzzle shapes. Here, we review findings related to the discovery of this novel auxin‐signaling mechanism at the cell surface. This signaling pathway is to a large extent independent of the well‐known Transport Inhibitor Response(TIR)–Auxin Signaling F‐Box(AFB) pathway, and instead requires Auxin Binding Protein 1(ABP1) interaction with the plasma membrane‐localized, transmembrane kinase(TMK) receptor‐like kinase to regulate ROP proteins. Once activated, ROP influences cytoskeletal organization and inhibits endocytosis of the auxin transporter PIN1. The present review focuses on ROP signaling and its self‐organizing feature allowing ROP proteins to serve as a bustling signal decoder and integrator for plant cell morphogenesis.     

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.     

Molecular interactions between light and hormone signaling to control plant growth

As sessile organisms, plants modulate their growth rate and development according to the continuous variation in the conditions of their surrounding environment, an ability referred to as plasticity. This ability relies on a web of interactions between signaling pathways triggered by endogenous and environmental cues. How changes in environmental factors are interpreted by the plant in terms of developmental or growth cues or, in other words, how they contribute to plant plasticity is a current, major question in plant biology. Light stands out among the environmental factors that shape plant development. Plants have evolved systems that allow them to monitor both quantitative and qualitative differences in the light that they perceive, that render important changes in their growth habit. In this review we focus on recent findings about how information from this environmental cue is integrated during de-etiolation and in the shade-avoidance syndrome, and modulated by several hormone pathways—the endogenous cues. In some cases the interaction between a hormone and the light signaling pathways is reciprocal, as is the case of the gibberellin pathway, whereas in other cases hormone pathways act downstream of the environmental cue to regulate growth. Moreover, the circadian clock adds an additional layer of regulation, which has been proposed to integrate the information provided by light with that provided by hormone pathways, to regulate daily growth.     

叶脉网络功能性状及其生态学意义

叶脉网络结构是叶脉系统在叶片里的分布和排列样式。早期叶脉网络结构研究主要集中在其分类学意义上; 近年来叶脉网络功能性状及其在植物水分利用上的意义已成为植物生态学研究的热点。该文介绍了叶脉网络功能性状的指标体系(包括叶脉密度、叶脉直径、叶脉之间的距离、叶脉闭合度等), 综述了叶脉网络功能性状与叶脉系统功能(包括水分、养分和光合产物等物质运输、机械支撑和虫害防御等)的关系, 叶脉网络功能性状与叶片其他功能性状(包括比叶重、叶寿命、光合速率、叶片大小、气孔密度等)的协同变异和权衡关系, 以及叶脉网络功能性状随环境因子(包括水分、温度、光照等)的变化规律等方面的最新研究进展。此外, 叶脉网络功能性状的研究成果也被应用于古环境重建、城市交通规划、流域规划及全球变化研究中。由于叶脉网络功能性状是环境因子与系统发育共同作用的结果, 未来开展分子—叶片—植物—生态系统等多尺度的叶脉网络功能性状研究, 理清叶脉网络功能性状与气孔失水—茎干导水—根系吸水等植物水分利用的关系, 将为预测植物及生态系统对全球变化的响应提供新的启示。     

叶甲科昆虫的定殖机制

在复杂的生态环境中,叶甲科昆虫利用寄主植物的挥发物作为嗅觉信号,并协同视觉信号共同作用以提高其远距离搜寻和定位寄主的效率。叶甲科昆虫也释放信息素来调节同种或者异种个体的行为反应;找到寄主植物后,叶子的形态学特性和化学组分的浓度等接触性因素就会影响叶甲科昆虫对寄主植物的最终选择。叶甲对上述这些信号物质的反应也受叶甲本身状态的影响,如生殖、滞育、饥饿、交配以及取食等。叶甲科昆虫对寄主植物的搜寻、辨识和接受、取食以及繁殖的过程受一种或多种因素协同影响。从嗅觉、视觉和触觉等方面对叶甲科昆虫的定殖机制做一综述。     

表观遗传调控植物叶片衰老的研究进展

植物叶片衰老是一个非常重要的发育过程,涉及大分子的有序分解从而将营养物质从叶片转移到其他器官,对植物的生存和适应至关重要。叶片衰老主要受植物的发育调控,但同时也受内部和外部环境因素的影响,涉及高度复杂的基因调控网络和多层级的调控。近年来的研究表明表观遗传是调控植物叶片衰老的一种重要调控方式。该研究综述了植物叶片衰老过程中的表观遗传调控机制,包括组蛋白修饰、DNA甲基化、ATP依赖的染色质重塑和非编码RNA介导的调控,并对该领域今后的发展方向进行了展望。     

彩叶植物叶片呈色分子机制研究进展

彩叶植物叶片呈现不同的颜色主要是受遗传因素和外部环境的共同作用,揭示彩叶植物叶片呈色机制对选育彩叶植物新品种和彩叶植物的应用推广具有重要理论和实践意义。目前对彩叶植物呈色机制的研究主要集中于叶片中色素变化、光合特性、叶片结构和环境条件等方面。该文主要対近年来有关彩叶植物叶片中叶绿素代谢途径、类胡萝卜素代谢途径、次生代谢途径、光合作用和叶绿体发育相关结构基因和转录因子调控机制的研究进展进行综述,并对以后的研究方向进行了展望,为培育彩叶植物新品种提供了理论基础,也为人工调控叶色以及叶色的定向遗传改良提供了参考。     

Of light and length: regulation of hypocotyl growth in Arabidopsis

At all stages, plant development results from a complex integration of multiple endogenous and environmental signals. The sedentary nature of plants strongly enhances the impact of the environment on plant development as compared to animal development. The embryonic and postembryonic seedling stem, called the hypocotyl, of the model species Arabidopsis (thale cress) has proved to be an excellent system for studying such signal interplay in the regulation of growth and developmental responses. The extension of the hypocotyl, which is regulated by a network of interacting factors, including light and plant hormones, is such a process. These regulatory factors often reciprocally regulate their biosynthesis and/or signalling. Here we present the current state of knowledge about the regulation of hypocotyl growth by a large repertoire of internal and external cues.     

Plasticity versus canalization: population differences in the timing of shade-avoidance responses

The reliability of environmental cues and costs of a fixed phenotype are two factors determining whether selection favors phenotypic plasticity or environmental specialization. This study examines the relationship between these two factors and the evolution of plant competitive strategies (plastic vs. fixed morphologies). In natural plant populations, shifts in light quality associated with foliar shade reliably indicate the presence of neighbors. These cues mediate plastic stem-elongation responses that often increase competitive ability and access to light. Using experimental light treatments (full sun, neutral shade, and foliar shade), genetic differences among populations of Abutilon theophrasti (velvetleaf) in average elongation and plasticity to foliar-shade cues were examined. Six populations, two from each of three site types (fields in continuous corn cultivation, fields undergoing corn-soy rotation, and weedy sites), were exposed to the light treatments at two stages in their life history. At the seedling stage, populations derived from cornfield sites exhibited higher, average elongation than populations from either rotating corn-soy fields or weedy areas. Because seedling elongation may delay shading of velvetleaf by corn, population differences may reflect adaptive responses to directional selection imposed by competitive conditions. However, the effects of simulated foliar shade on elongation were three times as great as the average population differences, and these comparatively higher levels of elongation were associated with an allocation cost. These results are consistent with the hypothesis that phenotypic plasticity may limit the evolution of specialists; reliable environmental cues enable individuals to facultatively adopt highly elongated, costly phenotypes in crowded patches while avoiding the costs of that phenotype in less crowded microsites. At later life-history stages, populations experiencing competition with corn exhibited lower plasticity to light quality than populations derived from weedy areas. Elongation at later nodes is maladaptive in cornfields because velvetleaf is ultimately incapable of overtopping corn; individuals that elongate therefore experience the cost of allocating to stems but fail to improve leaf exposure. The decreased responsiveness of cornfield populations to light quality is consistent with theoretical predictions in which reduced plasticity is favored when environmental cues fail to mediate an adaptive response.