植物根系形态对低磷胁迫应答的研究进展

本文综述了近年来植物对磷营养高效吸收有关的根系形态方面的研究进展, 总结了植物适应低磷胁迫的根系形态特征, 以及植物适应低磷胁迫根系形态变化的激素调控的内在机制, 着重阐述了植物适应低磷根系形态变化的分子生物学基础, 并对开展此类工作的有效途径进行了探讨。     

黄花苜蓿与蒺藜苜蓿对土壤低磷胁迫适应策略的比较研究

土壤有效磷(P)含量低是限制植物生长的主要因素之一。根形态变化和根系大量分泌以柠檬酸为主的有机酸是植物适应土壤P素缺乏的重要机制。以广泛分布于我国北方的重要豆科牧草黄花苜蓿(Medicago falcata)和豆科模式植物蒺藜苜蓿(M. truncatula)为材料, 采用砂培方法, 研究了低P胁迫对其植株生长、根系形态和柠檬酸分泌的影响, 对比了两种苜蓿适应低P胁迫的不同策略。结果表明: 1)低P处理显著抑制了蒺藜苜蓿与黄花苜蓿的地上部生长, 而对地下部生长影响较小, 从而导致根冠比增加。2)低P胁迫显著降低黄花苜蓿的总根长和侧根长, 而对蒺藜苜蓿的上述根系形态指标没有显著影响。3)低P胁迫促进两种苜蓿根系的柠檬酸分泌, 无论是在正常供P还是低P胁迫条件下, 黄花苜蓿根系分泌柠檬酸量显著高于蒺藜苜蓿根系。上述结果表明, 黄花苜蓿和蒺藜苜蓿对低P胁迫的适应策略不同, 低P胁迫下, 黄花苜蓿主要通过根系大量分泌柠檬酸, 活化根际难溶态P来提高对P的吸收, 而蒺藜苜蓿维持较大的根系是其适应低P胁迫的主要策略。     

植物miRNA在调节低磷胁迫响应中的作用

植物MicroRNA(miRNA)是一类内源性非编码小分子RNA,它们参与调节植物的生长、发育和代谢过程中多种基因的表达。近期的研究发现miRNA参与调节磷的吸收和利用,对植物适应低磷胁迫具有重要作用。本文概述了植物磷吸收和转运的机制,介绍了低磷胁迫下miRNA的表达水平变化,重点对miRNA在植物响应低磷胁迫中的作用,如改变根系结构、提高磷的转运和再利用效率、参与花青素和抗氧化物生物合成等进行了综述,以期为揭示植物低磷胁迫响应分子机制,提高植物对磷的吸收效率提供借鉴。     

磷高效利用杉木对低磷胁迫的适应性与内源激素的相关性

激素是植物适应逆境的重要信号物质, 从激素调控角度研究植物对养分匮乏环境的适应机制对磷高效营养基因型的选育具有重要意义。该研究通过分析被动忍受型(M1)与主动活化型(M4)两个磷高效利用杉木(Cunninghamia lanceolata)基因型在低磷胁迫下不同处理阶段的激素含量变化规律, 结合根系形态变化、干物质及养分分配规律, 研究磷高效利用杉木对低磷胁迫的适应性与内源激素的相关性。结果表明: 低磷处理下, 磷高效利用杉木M1与M4叶的激素含量与其适应特性之间无相关性, 而根系的激素含量与根系生长显著相关。低磷处理条件下, M1与M4根系中的IAA含量自27 h起表现为大于高磷对照, 且随时间延长呈增加趋势。根系中的IAA含量与根表面积、体积及根长等显著正相关, IAA的增加诱导了根系的增长, M1与M4均表现出一定的根系增长量。其中, M4存在明显的IAA由地上向基部积累的现象, M4的根系增生能力比M1更强。同时, 根系增长促使更多的干物质分配到根系, M4的根冠比在整个处理过程中均高于高磷对照。与IAA相同, M1与M4根系的ABA与GA₃含量总体也表现为低磷处理>高磷对照, 但随时间延长, 低磷条件下ABA与GA₃的含量呈下降趋势, 二者与根系增长量呈负相关关系。M1与M4根系内的ZT含量在低磷条件下也呈下降趋势, 且逐渐低于高磷对照, 而其与低磷适应特性间并无显著相关性。可见, 低磷胁迫下, 磷高效利用杉木M1与M4根系中的IAA、ABA与GA₃含量与其根系形态变化密切相关, 各器官的物质、能量、信息的综合调控是植物适应低磷逆境的重要生存策略。     

植物高效利用磷机制的研究进展

缺磷是限制目前农林业产量的一个重要因子,传统的农林业生产主要通过施肥和土壤改良来满足植物对磷的需求,近年来人们开始发掘磷高效利用植物来替代传统方法提高磷的利用效率。本文综述了国内外有关植物高效利用磷的形态学、生理学及遗传学作用机制,植物高效利用磷的机制主要包括：(1)磷高效利用植物能通过根系形态变化(包括根伸长、根轴变细、根毛数量和密度增大、侧根幼根数量增加及形成排根等)和根冠问的物质分配改变等形态学机制来适应磷胁迫;(2)在缺磷环境下磷高效利用植物能通过根系分泌物增加、菌根侵染、根系吸收动力学特征变化及植物磷素内循环加强等生理学机制来适应磷胁迫;(3)在磷亏缺的长期选择压力下,植物可通过某些“沉默”基因的诱导表达或DNA序列的特定形成相对稳定的磷营养遗传性状,由此通过遗传学机制来增加对土壤难溶态性磷的利用,使植物表现出较高的磷素利用效率。     

植物根系响应低磷胁迫的机理研究

磷是植物生长的必需营养元素之一。但大部分土壤中有效磷含量较低,难以满足植物生长的需求。作物磷效率遗传改良是解决土壤磷供应不足的有效途径。根系是植物吸收矿质营养元素的主要器官,其性状决定了植物对土壤磷的吸收利用效率。解析根系对低磷胁迫的响应机制是进行作物磷效率遗传改良的基础。主要介绍了近年来关于植物根系响应低磷胁迫机理的重要研究成果。     

菜豆根形态特性的基因型差异与磷效率

应用磷控释砂培以及计算机图象分析技术,研究了磷效率差异显的菜豆（Phaseolus vulgaris L.)亲本及其重组自交系后代的根形态特性及其与磷效率的关系。试验结果表明,供磷状况显影响菜豆根系形态学特性。在低磷胁迫下,菜豆根系总根长变短、根部生物量减少,根直径增大。菜豆根形态特性对低磷有效性的适应性反应具有显的基因型差异。在低磷条件下磷高效率基因型的根系比磷低效率基因型相对根部生物量较大、总根长较长,根表面积较大。异计分析表明,菜豆基根根形态特性在低磷条件下的适应性变化对磷效率的贡献远远大于主根,并且这些适应性变化是可以遗传的,表明通过对菜豆根形态特性进行遗传改良来提高磷效率有一定的可行性。     

在控制条件下云南松幼苗根系对低磷胁迫的响应

磷是控制生命过程的重要元素,植物在生长过程中需要大量的磷,低磷常导致一些植物发生适应性变化。云南松(PinusyunnanensisFranch.)以云南高原为起源和分布中心,其对低磷土壤环境表现出了很强的适应能力,广泛分布并正常生长于贫瘠的低磷红壤上,研究云南松对低磷环境的适应机制,对人类探索高效利用有限的磷素资源的方法具有现实意义。本实验通过对不同磷处理水平下培养的云南松幼苗根系生物量和根冠比等的研究,分析了云南松幼苗根系对低磷胁迫的响应。实验所用云南松种子采集自云南省通海县秀山森林公园内的健壮云南松林。结果表明:当磷浓度下降到0.5mmol/L时,云南松幼苗主根长度开始随磷浓度的降低而增加,根冠比随磷浓度的降低而增大,而侧根发生数没有随磷浓度的降低而显示出显著的增减规律,根系生物量也没有随磷浓度的降低而呈现出有规律的增减,根系生物量始终保持在一定的水平。进一步的分析表明:低磷胁迫下,云南松幼苗保证了物质分配对根的优先地位,以维持其根的生物量在一定水平,进而维持整个生命;云南松幼苗主要是靠主根长度的增加而不是靠侧根数量的增加来适应低磷环境。     

植物与低磷环境研究进展——诱导、适应与对策

自从20世纪70年代人们发现适应低磷土壤的作物根际磷的有效性明显增加的现象之后。植物与低磷环境的研究便引起了人们的重视。植物如何适应低磷环境和如何有效利用土壤磷素资源的问题已成为国内外当前的研究热点之一。研究表明,低磷条件下,植物根系形态结构会发生适应性变化,根冠间的物质分配会向根部倾斜使根冠比增加;植物根际酸度变化、有机酸分泌和磷酸酶释放有利于活化和利用土壤中的磷素资源;不同种类或品种的植物具有不同的磷营养效率基因型,具有不同亲和力的磷转运体,也具有不同的磷活化机制。人类对植物适应低磷机制的研究还将继续,揭示植物对低磷环境的响应对策和发掘植物有效利用磷素资源的潜力,在经济上和环保上均有非常现实的意义。     

高等植物对磷饥饿自我拯救的分子生物学机制

磷饥饿状态下,植物通过一系列生理、生化变化主动适应胁迫逆境,包括植物对土壤难溶性磷的活化、根系对低浓度有机磷的有效吸收,以及对吸收磷的再利用等。而这些生理生化反应都有其特定的分子生物学基础。本文着重综述与这三方面特性有关的分子生物学研究进展,包括与根系有机酸合成以活化难溶性磷有关的ＰＥＰ羧化酶（ＰＥＰＣ）;与有效吸收低浓度有机磷有关的高亲和力磷转运子;以及与利用生长介质中的有机磷有关的ＲＮａｓｅ、磷酸酶（ＡＰａｓｅ）;Ｃａ２＋－ＡＴＰａｓｅ;低磷营养胁迫导致的植物与菌根菌互作的分子生物学;以及磷饥饿诱导差异表达的基因等。     

植物根源逆境信使及其产生和传输

植物不仅仅是恶劣环境的被动受害者 ,而且它也具有对环境变化快速感知和主动适应的能力 ,逆境信息传递就是其适应不良环境的重要策略。本文主要介绍了环境胁迫下 ,调控植物地上部的根源信使、信使的产生和传输以及气孔运动对根源信使的响应机制等方面的研究进展。     

植物根源逆境信使及其产生和传输

植物不仅仅是恶劣环境的被动受害者,而且它也具有对环境变化快速感知和主动适应的能力,逆境信息传递就是其适应不良环境的重要策略。本文主要介绍了环境胁迫下,调控植物地上部的根源信使、信使的产生和传输以及气孔运动对根源信使的响应机制等方面的研究进展。     

Responses of root architecture development to low phosphorus availability: a review

Background

Phosphorus (P) is an essential element for plant growth and development but it is often a limiting nutrient in soils. Hence, P acquisition from soil by plant roots is a subject of considerable interest in agriculture, ecology and plant root biology. Root architecture, with its shape and structured development, can be considered as an evolutionary response to scarcity of resources.

Scope

This review discusses the significance of root architecture development in response to low P availability and its beneficial effects on alleviation of P stress. It also focuses on recent progress in unravelling cellular, physiological and molecular mechanisms in root developmental adaptation to P starvation. The progress in a more detailed understanding of these mechanisms might be used for developing strategies that build upon the observed explorative behaviour of plant roots.

Conclusions

The role of root architecture in alleviation of P stress is well documented. However, this paper describes how plants adjust their root architecture to low-P conditions through inhibition of primary root growth, promotion of lateral root growth, enhancement of root hair development and cluster root formation, which all promote P acquisition by plants. The mechanisms for activating alterations in root architecture in response to P deprivation depend on changes in the localized P concentration, and transport of or sensitivity to growth regulators such as sugars, auxins, ethylene, cytokinins, nitric oxide (NO), reactive oxygen species (ROS) and abscisic acid (ABA). In the process, many genes are activated, which in turn trigger changes in molecular, physiological and cellular processes. As a result, root architecture is modified, allowing plants to adapt effectively to the low-P environment. This review provides a framework for understanding how P deficiency alters root architecture, with a focus on integrated physiological and molecular signalling.     

Genetic improvement for phosphorus efficiency in soybean: a radical approach

Background

Low phosphorus (P) availability is a major constraint to soybean growth and production. Developing P-efficient soybean varieties that can efficiently utilize native P and added P in the soils would be a sustainable and economical approach to soybean production.

Scope

This review summarizes the possible mechanisms for P efficiency and genetic strategies to improve P efficiency in soybean with examples from several case studies. It also highlights potential obstacles and depicts future perspectives in ‘root breeding’.

Conclusions

This review provides new insights into the mechanisms of P efficiency and breeding strategies for this trait in soybean. Root biology is a new frontier of plant biology. Substantial efforts are now focusing on increasing soybean P efficiency through ‘root breeding’. To advance this area, additional collaborations between plant breeders and physiologists, as well as applied and theoretical research are needed to develop more soybean varieties with enhanced P efficiency through root modification, which might contribute to reduced use of P fertilizers, expanding agriculture on low-P soils, and achieving more sustainable agriculture.     

Ethylene and plant responses to nutritional stress

Although ethylene is known to be involved in plant response to a number of biotic and abiotic stresses, relatively little is known concerning its role in nutritional stress arising from nutrient deficiency or mineral toxicity. There is clear evidence for involvement of ethylene in the symbiosis between Rhizobium and legumes, and in the 'Strategy 1' response to Fe deficiency. Ethylene may also be generated during tissue necrosis induced by severe toxicities and deficiencies. Metal toxicity may generate ethylene through oxidative stress. Evidence for a more general role for ethylene in regulating plant responses to macronutrient deficiency is suggestive but incomplete. Few studies have addressed this interaction, and most published reports are difficult to interpret because of the unrealistic way that nutrient treatments were imposed. Deficiency of N and P appear to interact with ethylene production and sensitivity. A role for ethylene in mediating adaptive responses to P stress is suggested by the fact that P stress can induce a variety of morphological changes in root systems that are also affected by ethylene, such as gravitropism, aerenchyma formation, and root hair development. Other adaptive responses include senescence or abscission of plant parts which cannot be supported by the plant. Ethylene and other plant hormones may be involved in mediating the stress signal to generate these responses. Although existing literature is inconclusive, we speculate that ethylene may play an important role in mediating the morphological and physiological plasticity of plant responses to nutrient patches in time and space, and especially root responses to P stress.     

Root colonization and iron nutritional status of a Pseudomonas fluorescens in different plant species

Root colonization and induction of an iron stress regulated promoter for siderophore production by Pseudomonas fluorescens 2-79RLI was studied in vitro and in the rhizosphere of different plant species. P. fluorescens 2-79RLI was previously genetically modified with an iron regulated ice nucleation reporter, which allowed calibration of ice nucleation activity with siderophore production. Initial experiments examined ice nucleation activity and siderophore production under different growth conditions in vitro. These studies demonstrated that P. fluorescens 2-79RLI could utilize both Fe-citrate and Fe-phytosiderophore as iron sources, suggesting that production of these compounds by plants would increase iron availability for P. fluorescens 2-79RLI in the rhizosphere. Fe demand and Fe stress were further shown to be a function of nutrient availability and were reduced when carbon was limiting for growth. Subsequent experiments extended these observations to rhizosphere cells. Cells were sampled from the rhizosphere and the rhizoplane. Results of a soil microcosm experiment showed that Fe stress was reduced for P. fluorescens 2-79RLI in the barley rhizosphere as compared to the cells in the rhizosphere.of lupin. In lupin, relative Fe stress of P. fluorescens 2-79RLI was greater at the root tip than in the lateral root zone. In a second experiment comparing zucchini and bean, iron stress was greater for P. fluorescens 2-79RLI associated with zucchini than with bean. In a third experiment with rape plants under P deficient conditions, addition of soluble P was shown to increase Fe stress for P. fluorescens 2-79RLI located at the root tip, but not in the lateral root zone. This study showed that Fe stress of P. fluorescens 2-79RLI in the rhizosphere may be influenced by plant species, P source, root zone and localization of the cells within the rhizosphere.     

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。     

Auxin: a master regulator in plant root development

The demand for increased crop productivity and the predicted challenges related to plant survival under adverse environmental conditions have renewed the interest in research in root biology. Various physiological and genetic studies have provided ample evidence in support of the role of plant growth regulators in root development. The biosynthesis and transport of auxin and its signaling play a crucial role in controlling root growth and development. The univocal role of auxin in root development has established it as a master regulator. Other plant hormones, such as cytokinins, brassinosteroids, ethylene, abscisic acid, gibberellins, jasmonic acid, polyamines and strigolactones interact either synergistically or antagonistically with auxin to trigger cascades of events leading to root morphogenesis and development. In recent years, the availability of biological resources, development of modern tools and experimental approaches have led to the advancement of knowledge in root development. Research in the areas of hormone signal perception, understanding network of events involved in hormone action and the transport of plant hormones has added a new dimension to root biology. The present review highlights some of the important conceptual developments in the interplay of auxin and other plant hormones and associated downstream events affecting root development.