植物向重力反应中PIN-FORMED介导的生长素极性运输调控

植物的向性,即植物对光或重力等环境刺激信号产生的定向生长反应。在向重力性反应中,植物器官将重力感知为定向环境信号,来控制其器官的生长方向以促进生存。植物激素生长素及其极性运输在植物向重力反应中起着决定性的调控作用。质膜定位的生长素输出蛋白PIN-FORMED(PIN)通过动态的亚细胞极性定位,改变生长素运输的方向以响应环境刺激,由此植物器官间建立的生长素浓度梯度是细胞差异化伸长和器官弯曲的基础,来调控植物的形态建成和生长发育过程。本文主要讨论发生在植物重力感受细胞内早期重力感知和信号转导机制的最新研究进展、PIN介导的生长素极性运输、PIN的极性定位以及质膜蛋白丰度的调控机制等。     

生长素调控根向重力性在植物激素实验教学中的应用

根是植物重要的器官,其向重力生长是由生长素的浓度与分布所调控的,是植物对环境信号作出的生理反应。本文对模式植物拟南芥的根进行改变重力方向的刺激,借助含有响应生长素的分子标记的DR5rev:GFP、DR5:3xVenus和DR5:GUS转基因株系,可视化在重力方向改变刺激下根尖两侧的生长素不对称分布,同时观察生长素极性运输载体的相关突变体pin2-T、aux1-T在重力方向改变刺激下的表型,帮助学生深入理解生长素调控拟南芥根向重力生长的生理机制。     

植物非经典生长素信号转导通路解析

植物激素生长素参与调控植物生长发育的各个过程,包括胚胎发育、器官发生和向性运动等。植物通过协调生长素的合成代谢、极性运输以及信号转导来实现对不同生长发育过程的精准调控。生长素的功能依赖于其信号被感知后经由信号转导通路转换为下游复杂多样的反应。经典的生长素信号转导通路阐明了细胞核内从SCF~(TIR1/AFB)受体到Aux/IAA蛋白的泛素化降解最终通过ARF转录因子调控基因转录的完整生长素响应过程。该核内信号通路揭示了生长素转录调控生长发育的诸多分子机制,但植物生长发育调控过程中仍有许多生长素响应过程无法通过该经典信号通路解析。重点阐述生长素非经典信号通路的调控机制及其对植物生长发育的重要作用,并讨论和展望生长素非经典信号通路研究目前所面临的挑战以及研究前景。     

生长素介导环境信号调控植物的生长发育

植物是一类营固着生活的自养型生物, 如何更好地适应周围环境对植物的生存至关重要。生长素是调控植物生长发育的重要激素之一。近年来的研究发现, 生长素不仅能够响应内在的发育信号, 而且能够介导各种环境信号, 参与植物生长发育和生长反应的调控。该文主要从光信号、温度信号、重力信号、营养元素和金属离子信号等方面重点阐述生长素如何介导上述各种不同的环境信号, 从而调控植物的生长发育。     

生长素调控植物重力反应的分子机理研究

重力反应是植物对环境的一种适应现象。生长素参与植物环境适应与发育调控的过程,重力反应过程的核心之一是在重力反应器官形成生长素的浓度梯度,诱导下游基因的差异表达。生长素的合成、代谢、极性运输及信号转导在此过程中发挥了关键作用。该文以拟南芥和水稻的研究为基础,综述了近几年对生长素调控植株重力反应的分子机理的研究进展,并对该领域未来的研究进行展望。     

封面说明

<正>独脚金内酯(strigolactones,SLs)是一类新型植物激素,能够抑制植物分枝的生长发育。近年来,关于SLs合成与信号在调控水稻株型方面的研究取得了重要进展。研究发现,独脚金内酯不仅可以调控水稻的分蘖数目,而且可影响分蘖角度和株高,进而影响穗部形态和籽粒大小,对水稻的产量具有显著的影响。鉴于独脚金内酯对于水稻株型的综合调控功     

生长素信号对植物侧根发育的调控

植物根系是汲取水分、营养的重要器官,而侧根是植物根系重要的组成部分。生长素是调控侧根生长发育的核心因子。该文综述了生长素信号在直根系模式植物拟南芥以及须根系模式作物水稻中侧根发育调控中的研究进展,对生长素信号调控侧根起始模型、Aux/IAA介导的生长素信号对植物侧根发育调控这两个方面进行了阐述,并对拟南芥与水稻的侧根发育进行比较,最后对该研究领域进行了展望。     

柳暗花明：胞外生长素信号感受的新突破

生长素在植物生长发育过程中发挥重要作用,其信号转导机制一直是植物学领域关注的热点。前期研究表明,ABP1-TMK分子模块参与胞外生长素信号感受,但ABP1作为生长素受体备受争议。近期,福建农林大学徐通达团队和杨贞标团队鉴定到ABL蛋白作为生长素结合蛋白参与胞外生长素信号感受。与传统的功能冗余不同, ABL和ABP1通过蛋白结构的相似性实现功能补偿效应,进而与TMK在细胞膜上形成复合体,作为胞外生长素的共受体介导生长素信号驱动的快速反应。该研究深入解析了胞外生长素信号感受的重要机制,是生长素研究领域的重大突破。     

糖在植物中的感知与信号传导研究进展

糖在植物中不仅用于能量的代谢,同时还作为信号分子调控植物的生长和发育.本文对植物体内糖信号的产生、糖类对植物体生长发育与胁迫反应的调节作用、糖与植物激素信号之间的传感关系以及糖信号调节的分子机制等的研究进展进行了综述.     

生长素与乙烯信号途径及其相互关系研究进展

长期的研究表明,生长素在调节植物生长发育的各种生理活动中起关键作用,但对它如何调控这些生理活动却缺乏系统和深入的了解。最近,细胞核内生长素信号途径的发现为揭示其作用机制带来了曙光。乙烯参与果实成熟及植物对逆境的反应等生理活动,其信号途径也已得到部分阐明。越来越多的证据表明,乙烯的作用与生长素对植物生长发育的调控之间有密切的联系。该文概述了生长素与乙烯信号途径的研究进展及其相互关系,讨论了生长素在植物三重反应中的作用;并对生长素与乙烯相互关系研究中存在的问题及研究前景进行了探讨。     

LAZY3 interacts with LAZY2 to regulate tiller angle by modulating shoot gravity perception in rice

Starch biosynthesis in gravity-sensing tissues of rice shoot determines the magnitude of rice shoot gravitropism and thus tiller angle. However, the molecular mechanism underlying starch biosynthesis in rice gravity-sensing tissues is still unclear. We characterized a novel tiller angle gene LAZY3 (LA3) in rice through map-based cloning. Biochemical, molecular and genetic studies further demonstrated the essential roles of LA3 in gravity perception of rice shoot and tiller angle control. The shoot gravitropism and lateral auxin transport were defective in la3 mutant upon gravistimulation. We showed that LA3 encodes a chloroplast-localized tryptophan-rich protein associated with starch granules via Tryptophan-rich region (TRR) domain. Moreover, LA3 could interact with the starch biosynthesis regulator LA2, determining starch granule formation in shoot gravity-sensing tissues. LA3 and LA2 negatively regulate tiller angle in the same pathway acting upstream of LA1 to mediate asymmetric distribution of auxin. Our study defined LA3 as an indispensable factor of starch biosynthesis in rice gravity-sensing tissues that greatly broadens current understanding in the molecular mechanisms underlying the starch granule formation in gravity-sensing tissues, and provides new insights into the regulatory mechanism of shoot gravitropism and rice tiller angle.     

TAC4 controls tiller angle by regulating the endogenous auxin content and distribution in rice

Tiller angle, an important component of plant architecture, greatly influences the grain yield of rice (Oryza sativa L.). Here, we identified Tiller Angle Control 4 (TAC4) as a novel regulator of rice tiller angle. TAC4 encodes a plant‐specific, highly conserved nuclear protein. The loss of TAC4 function leads to a significant increase in the tiller angle. TAC4 can regulate rice shoot gravitropism by increasing the indole acetic acid content and affecting the auxin distribution. A sequence analysis revealed that TAC4 has undergone a bottleneck and become fixed in indica cultivars during domestication and improvement. Our findings facilitate an increased understanding of the regulatory mechanisms of tiller angle and also provide a potential gene resource for the improvement of rice plant architecture.     

LAZY1 controls rice shoot gravitropism through regulating polar auxin transport

Tiller angle of rice （Oryza sativa L.） is an important agronomic trait that contributes to grain production, and has long attracted attentions of breeders for achieving ideal plant architecture to improve grain yield. Although enormous efforts have been made over the past decades to study mutants with extremely spreading or compact tillers, the molecular mechanism underlying the control of tiller angle of cereal crops remains unknown. Here we report the cloning of the LAZY1 （LA1） gene that regulates shoot gravitropism by which the rice tiller angle is controlled. We show that LA1, a novel grass-specific gene, is temporally and spatially expressed, and plays a negative role in polar auxin transport （PAT）. Loss-of-function of LA1 enhances PAT greatly and thus alters the endogenous IAA distribution in shoots, leading to the reduced gravitropism, and therefore the tiller-spreading phenotype of rice plants.     

Understanding the Regulatory Mechanisms of Rice Tiller Angle,Then and Now

Rice is one of the most important crops worldwide, whose yield is vital to human nutrition in the context of a rapidly growing world population. Plant architecture significantly affects grain yield, which is to a large extent determined by tiller angle and tiller number. Tiller angle is the angle between the primary tiller and the main culm. Its regulation is complex and is influenced by multiple environmental and genetic factors. This review provides an overview of the regulation of tiller angle in rice, with particular focus on the roles of the growth environment and method of cultivation; phytohormones such as auxin, gibberellins, and strigolactones; gravity; and genes related to the control of tiller angle. The major research foci and the outlook for research into the regulation of tiller angle in rice are discussed.

     

Over-expression of OsPIN2 leads to increased tiller numbers, angle and shorter plant height through suppression of OsLAZY1

Crop architecture parameters such as tiller number, angle and plant height are important agronomic traits that have been considered for breeding programmes. Auxin distribution within the plant has long been recognized to alter architecture. The rice (Oryza sativa L.) genome contains 12 putative PIN genes encoding auxin efflux transporters, including four PIN1 and one PIN2 genes. Here, we report that over-expression of OsPIN2 through a transgenic approach in rice (Japonica cv. Nipponbare) led to a shorter plant height, more tillers and a larger tiller angle when compared with wild type (WT). The expression patterns of the auxin reporter DR5::GUS and quantification of auxin distribution showed that OsPIN2 over-expression increased auxin transport from the shoot to the root-shoot junction, resulting in a non-tissue-specific accumulation of more free auxin at the root-shoot junction relative to WT. Over-expression of OsPIN2 enhanced auxin transport from shoots to roots, but did not alter the polar auxin pattern in the roots. Transgenic plants were less sensitive to N-1-naphthylphthalamic acid, an auxin transport inhibitor, than WT in their root growth. OsPIN2-over-expressing plants had suppressed the expression of a gravitropism-related gene OsLazy1 in the shoots, but unaltered expression of OsPIN1b and OsTAC1, which were reported as tiller angle controllers in rice. The data suggest that OsPIN2 has a distinct auxin-dependent regulation pathway together with OsPIN1b and OsTAC1 controlling rice shoot architecture. Altering OsPIN2 expression by genetic transformation can be directly used for modifying rice architecture.     

OsmiR167a‐targeted auxin response factors modulate tiller angle via fine‐tuning auxin distribution in rice

Rice tiller angle determines plant growth density and further contributes grain production. Although a few genes have been characterized to regulate tiller angle in rice, the molecular mechanism underlying the control of tiller angle via microRNA is poorly understood. Here, we report that rice tiller angle is controlled by OsmiR167a‐targeted auxin response factors OsARF12, OsARF17 and OsARF25. In the overexpression of OsMIR167a plants, the expression of OsARF12, OsARF17 and OsARF25 was severely repressed and displayed larger tiller angle as well as the osarf12/osarf17 and osarf12/ osarf25 plants. In addition, those plants showed compromised abnormal auxin distribution and less sensitive to gravity. We also demonstrate that OsARF12, OsARF17 and OsARF25 function redundantly and might be involved in HSFA2D and LAZY1‐dependent asymmetric auxin distribution pathway to control rice tiller angle. Our results reveal that OsmiR167a represses its targets, OsARF12, OsARF17 and OsARF25, to control rice tiller angle by fine‐tuning auxin asymmetric distribution in shoots.     

Gravity signal transduction in primary roots

AIMS: The molecular mechanisms that correlate with gravity perception and signal transduction in the tip of angiosperm primary roots are discussed. SCOPE: Gravity provides a cue for downward orientation of plant roots, allowing anchorage of the plant and uptake of the water and nutrients needed for growth and development. Root gravitropism involves a succession of physiological steps: gravity perception and signal transduction (mainly mediated by the columella cells of the root cap); signal transmission to the elongation zone; and curvature response. Interesting new insights into gravity perception and signal transduction within the root tip have accumulated recently by use of a wide range of experimental approaches in physiology, biochemistry, genetics, genomics, proteomics and cell biology. The data suggest a network of signal transduction pathways leading to a lateral redistribution of auxin across the root cap and a possible involvement of cytokinin in initial phases of gravicurvature. CONCLUSION: These new discoveries illustrate the complexity of a highly redundant gravity-signalling process in roots, and help to elucidate the global mechanisms that govern auxin transport and morphogenetic regulation in roots.     

Auxin transport and gravitational research: perspectives

Gravity is a fundamental factor which affects all living organisms. Plant development is well adapted to gravity by directing roots downward and shoots upwards. For more than a century, plant biologists have been fascinated to describe the molecular mechanisms underlying the gravitropic response of plants. Important progress towards signal perception, transduction, and response has been made, but new tools are beginning to uncover the regulatory networks for gravitropic control. We summarise recent progress in study of gravitropism and discuss strategies to identify the molecular basis of the gravity response in Arabidopsis thaliana. This will put us on a road towards the molecular systems biology of the Arabidopsis gravitropic response.