植物MPK信号通路与信号分子H2O2和NO之间的相互关系

促分裂原活化蛋白激酶(MPK)级联途径是真核细胞中普遍存在且保守的信号传导通路,广泛参与植物生长发育和植物抵抗生物和非生物胁迫的防御反应。过氧化氢(H2O2)和一氧化氮(NO)作为重要的信使分子也广泛参与植物生长发育和防御反应的信号传导。近年来,研究也表明MPK信号通路与信号分子H2O2和NO之间存在着多种复杂的关系。一方面,在一些刺激的信号传递过程中,MPK信号通路参与了信号分子H2O2和NO的产生、清除或其信号的向下传递等过程;另一方面,在有些刺激的信号传递过程中,它们位于不同的信号传递途经中,行使不同的功能。本文就目前植物MPK信号通路与H2O2和NO之间相互关系的研究现状进行了综述和分析,并指出了该研究领域存在的问题。     

Role of Pathogen-Related Protein 10 (PR 10) under Abiotic and Biotic Stresses in Plants

Members of the Pathogenesis Related (PR) 10 protein family have been identified in a variety of plant species and a wide range of functions ranging from defense to growth and development has been attributed to them. PR10 protein possesses ribonuclease (RNase) activity, interacts with phytohormones, involved in hormone-mediated signalling, afforded protection against various phytopathogenic fungi, bacteria, and viruses particularly in response to biotic and abiotic stresses. The resistance mechanism of PR10 protein may include activation of defense signalling pathways through possible interacting proteins involved in mediating responses to pathogens, degradation of RNA of the invading pathogens. Moreover, several morphological changes have been shown to accompany the enhanced abiotic stress tolerance. In this review, the possible mechanism of action of PR10 protein against biotic and abiotic stress has been discussed. Furthermore, our findings also confirmed that the in vivo Nitric oxide (NO) is essential for most of environmental abiotic stresses and disease resistance against pathogen infection. The proper level of NO may be necessary and beneficial, not only in plant response to the environmental abiotic stress, but also to biotic stress. The updated information on this interesting group of proteins will be useful in future research to develop multiple stress tolerance in plants.     

Light perception in plant disease defence signalling

Light is a predominant factor in the control of plant growth, development and stress responses. Many biotic stress responses in plants are therefore specifically adjusted by the prevailing light conditions. The plant cell is equipped with sophisticated light-sensing mechanisms that are localised inside and outside of the chloroplast and the nucleus. Recent progress has provided models of how the signalling pathways that are involved in light perception and in defence could operate and interact to form a plant defence network. Such a signalling network includes systems to sense light and regulate gene expression. Photo-produced H(2)O(2) and other reactive oxygen species in the cell also play an essential role in this regulatory network, controlling biotic and abiotic stress responses.     

Rapid transmission of oxidative and nitrosative stress signals from roots to shoots in Arabidopsis.

Protein kinases play a central role in signal transduction pathways in eukaryotes. A highly conserved group of kinases, termed mitogen-activated-protein kinases (MAPKs) was shown to mediate many diverse stress responses. In plants, MAPKs were shown to function in resistance responses to many biotic and abiotic stresses. Here, we show that exposure of Arabidopsis roots to hydrogen peroxide or to nitric oxide resulted in rapid activation of protein kinases in the shoots that exhibited MAPK properties. The same pattern of kinases was induced by direct injection of these compounds into leaves, indicating accurate long-distance transmission of H2O2 and NO signals. These results are important for the understanding of redox signal transmission from the rhizosphere throughout the plant.     

Polyamines, polyamine oxidases and nitric oxide in development, abiotic and biotic stresses

Nitric oxide (NO), polyamines (PAs), diamine oxidases (DAO) and polyamine oxidases (PAO) play important roles in wide spectrum of physiological processes such as germination, root development, flowering and senescence and in defence responses against abiotic and biotic stress conditions. This functional overlapping suggests interaction of NO and PA in signalling cascades. Exogenous application of PAs putrescine, spermidine and spermine to Arabidopsis seedlings induced NO production as observed by fluorimetry and fluorescence microscopy using the NO-binding fluorophores DAF-2 and DAR-4M. The observed NO release induced by 1 mM spermine treatment in the Arabidopsis seedlings was very rapid without apparent lag phase. These observations pave a new insight into PA-mediated signalling and NO as a potential mediator of PA actions. When comparing the functions of NO and PA in plant development and abiotic and biotic stresses common to both signalling components it can be speculated that NO may be a link between PA-mediated stress responses filing a gap between many known physiological effects of PAs and amelioration of stresses. NO production indicated by PAs could be mediated either by H₂O₂, one reaction product of oxidation of PAs by DAO and PAO, or by unknown mechanisms involving PAs, DAO and PAO.     

Hydrogen peroxide-induced gene expression in Arabidopsis thaliana

Hydrogen peroxide (H(2)O(2)) is generated in plants after exposure to a variety of biotic and abiotic stresses, and has been shown to induce a number of cellular responses. Previously, we showed that H(2)O(2) generated during plant-elicitor interactions acts as a signaling molecule to induce the expression of defense genes and initiate programmed cell death in Arabidopsis thaliana suspension cultures. Here, we report for the first time the identification by RNA differential display of four genes whose expression is induced by H(2)O(2). These include genes that have sequence homology to previously identified Arabidopsis genes encoding a late embryogenesis-abundant protein, a DNA-damage repair protein, and a serine/threonine kinase. Their putative roles in H(2)O(2)-induced defense responses are discussed.     

NO是植物应激反应的信号分子

根据NO的性质和可能的产生途径,略述了生物胁迫(病原菌侵害)和干旱胁迫、盐胁迫、极端温度、机械损伤、臭氧和紫外辐射等各种非生物胁迫信号与NO信号分子的偶联及其信号的级联途径,概括了NO可能介导的生物过程,讨论了NO通过其下游信号过程对与细胞的生理影响以及该下游信号过程所涉及到的cGMP、cADPR的产生和NO与其它信号分子(ROS、SA、ABA等)的协同作用,表明胁迫诱导的NO爆发是激发、启动和装备植物细胞的重要信号级联环节,这个环节能使植物细胞处于应激状态,并迅速作出反应,形成一系列适应机制。     

Involvement of hydrogen peroxide and nitric oxide in salt resistance in the calluses from Populus euphratica

Nitric oxide (NO) and hydrogen peroxide (H2O2) function as signalling molecules in plants under abiotic and biotic stresses. Calluses from Populus euphratica, which show salt tolerance, were used to study the interaction of NO and H2O2 in plant adaptation to salt resistance. The nitric oxide synthase (NOS) activity was identified in the calluses, and this activity was induced under 150 mM NaCl treatment. Under 150 mM NaCl treatment, the sodium (Na) percentage decreased, but the potassium (K) percentage and the K/Na ratio increased in P. euphratica calluses. Application of glucose/glucose oxidase (G/GO, a H2O2 donor) and sodium nitroprusside (SNP, a NO donor) revealed that both H2O2 and NO resulted in increased K/Na ratio in a concentration-dependent manner. Diphenylene iodonium (DPI, an NADPH oxidase inhibitor) counteracted H2O2 and NO effect by increasing the Na percentage, decreasing the K percentage and K/Na ratio. NG-monomethyl-L-Arg monoacetate (NMMA, an NO synthase inhibitor) and 2-phenyl-4,4,5,5-tetramethyl-imidazoline-1-oxyl-3-oxyde (PTIO, a specific NO scavenger) only reversed NO effect, but did not block H2O2 effect. The increased activity of plasma membrane (PM) H+ -ATPase caused by salt stress was reversed by treatment with DPI and NMMA. Exogenous H2O2 increased the activity of PM H+ -ATPase, but the effect could not be diminished by NMMA and PTIO. The NO-induced increase of PM H+ -ATPase can be reversed by NMMA and PTIO, but not by DPI. Western blot analysis demonstrated that NO and H2O2 stimulated the expression of PM H+ -ATPase in P. euphratica calluses. These results indicate that NO and H2O2 served as intermediate molecules in inducing salt resistance in the calluses from P. euphratica under slat stress by increasing the K/Na ratio, which was dependent on the increased PM H+ -ATPase activity.     

The polyamines and their catabolic products are significant players in the turnover of nitrogenous molecules in plants

Polyamines (PAs) are nitrogenous molecules which play a well-established role in most cellular processes during growth and development under physiological or biotic/abiotic stress conditions. The molecular mode(s) of PA action have only recently started to be unveiled, and comprehensive models for their molecular interactions have been proposed. Their multiple roles are exerted, at least partially, through signalling by hydrogen peroxide (H(2)O(2)), which is generated by the oxidation/back-conversion of PAs by copper amine oxidases and PA oxidases. Accumulating evidence suggests that in plants the cellular titres of PAs are affected by other nitrogenous compounds. Here, we discuss the state of the art on the possible nitrogen flow in PAs, their interconnection with nitrogen metabolism, as well as the signalling roles of PA-derived H(2)O(2) during some developmental processes and stress responses.     

植物细胞活性氧种类、代谢及其信号转导

越来越明显的证据表明,植物体十分活跃的产生着活性氧并将之作为信号分子、进而控制着诸如细胞程序性死亡、非生物胁迫响应、病原体防御和系统信号等生命过程,而不仅是传统意义上的活性氧是有氧代谢的附产物。日益增多的证据显示,由脱落酸、水杨酸、茉莉酸与乙烯以及活性氧所调节的激素信号途径,在生物和非生物胁迫信号的“交谈”中起重要作用。活性氧最初被认为是动物吞噬细胞在宿主防御反应时所释放的附产物,现在的研究清楚的表明,活性氧在动物和植物细胞信号途径中均起作用。活性氧可以诱导细胞程序性死亡或坏死、可以诱导或抑制许多基因的表达,也可以激活上述级联信号。近来生物化学与遗传学研究证实过氧化氢是介导植物生物胁迫与非生物胁迫的信号分子,过氧化氢的合成与作用似乎与一氧化氮有关系。过氧化氢所调节的下游信号包括钙“动员”、蛋白磷酸化和基因表达等。     

Modulation of nitric oxide bioactivity by plant haemoglobins

Nitric oxide (NO) is a highly reactive signalling molecule that has numerous targets in plants. Both enzymatic and non-enzymatic synthesis of NO has been detected in several plant species, and NO functions have been characterized during diverse physiological processes such as plant growth, development, and resistance to biotic and abiotic stresses. This wide variety of effects reflects the basic signalling mechanisms that are utilized by virtually all mammalian and plant cells and suggests the necessity of detoxification mechanisms to control the level and functions of NO. During the last two years an increasing number of reports have implicated non-symbiotic haemoglobins as the key enzymatic system for NO scavenging in plants, indicating that the primordial function of haemoglobins may well be to protect against nitrosative stress and to modulate NO signalling functions. The biological relevance of plant haemoglobins during specific conditions of plant growth and stress, and the existence of further enzymatic and non-enzymatic NO scavenging systems, suggest the existence of precise NO modulation mechanisms in plants, as observed for different NO sources.     

Hydrogen Peroxide in Plants： a Versatile Molecule of the Reactive Oxygen Species Network

Plants often face the challenge of severe environmental conditions, which include various biotic and abiotic stresses that exert adverse effects on plant growth and development. During evolution, plants have evolved complex regulatory mechanisms to adapt to various environmental stressors. One of the consequences of stress is an increase in the cellular concentration of reactive oxygen species （ROS）, which are subsequently converted to hydrogen peroxide （H2O2）. Even under normal conditions, higher plants produce ROS during metabolic processes. Excess concentrations of ROS result in oxidative damage to or the apoptotic death of cells. Development of an antioxidant defense system in plants protects them against oxidative stress damage. These ROS and, more particularly, H2O2, play versatile roles in normal plant physiological processes and in resistance to stresses. Recently, H2O2 has been regarded as a signaling molecule and regulator of the expression of some genes in cells. This review describes various aspects of H2O2 function, generation and scavenging, gene regulation and cross-links with other physiological molecules during plant growth, development and resistance responses.     

Hydrogen peroxide signalling

Recent biochemical and genetic studies confirm that hydrogen peroxide is a signalling molecule in plants that mediates responses to abiotic and biotic stresses. Signalling roles for hydrogen peroxide during abscisic-acid-mediated stomatal closure, auxin-regulated root gravitropism and tolerance of oxygen deprivation are now evident. The synthesis and action of hydrogen peroxide appear to be linked to those of nitric oxide. Downstream signalling events that are modulated by hydrogen peroxide include calcium mobilisation, protein phosphorylation and gene expression. Calcium and Rop signalling contribute to the maintenance of hydrogen peroxide homeostasis.     

S-Nitrosylated proteins in pea (Pisum sativum L.) leaf peroxisomes: changes under abiotic stress

Peroxisomes, single-membrane-bounded organelles with essentially oxidative metabolism, are key in plant responses to abiotic and biotic stresses. Recently, the presence of nitric oxide (NO) described in peroxisomes opened the possibility of new cellular functions, as NO regulates diverse biological processes by directly modifying proteins. However, this mechanism has not yet been analysed in peroxisomes. This study assessed the presence of S-nitrosylation in pea-leaf peroxisomes, purified S-nitrosylated peroxisome proteins by immunoprecipitation, and identified the purified proteins by two different mass-spectrometry techniques (matrix-assisted laser desorption/ionization tandem time-of-flight and two-dimensional nano-liquid chromatography coupled to ion-trap tandem mass spectrometry). Six peroxisomal proteins were identified as putative targets of S-nitrosylation involved in photorespiration, β-oxidation, and reactive oxygen species detoxification. The activity of three of these proteins (catalase, glycolate oxidase, and malate dehydrogenase) is inhibited by NO donors. NO metabolism/S-nitrosylation and peroxisomes were analysed under two different types of abiotic stress, i.e. cadmium and 2,4-dichlorophenoxy acetic acid (2,4-D). Both types of stress reduced NO production in pea plants, and an increase in S-nitrosylation was observed in pea extracts under 2,4-D treatment while no total changes were observed in peroxisomes. However, the S-nitrosylation levels of catalase and glycolate oxidase changed under cadmium and 2,4-D treatments, suggesting that this post-translational modification could be involved in the regulation of H(2)O(2) level under abiotic stress.     

Revealing on hydrogen sulfide and nitric oxide signals co-ordination for plant growth under stress conditions

In the recent times, plants are facing certain types of environmental stresses, which give rise to formation of reactive oxygen species (ROS) such as hydroxyl radicals, hydrogen peroxides, superoxide anions and so on. These are required by the plants at low concentrations for signal transduction and at high concentrations, they repress plant root growth. Apart from the ROS activities, hydrogen sulfide (H₂S) and nitric oxide (NO) have major contributions in regulating growth and developmental processes in plants, as they also play key roles as signaling molecules and act as chief plant immune defense mechanisms against various biotic as well as abiotic stresses. H₂S and NO are the two pivotal gaseous messengers involved in growth, germination and improved tolerance in plants under stressed and non-stress conditions. H₂S and NO mediate cell signaling in plants as a response to several abiotic stresses like temperature, heavy metal exposure, water and salinity. They alter gene expression levels to induce the synthesis of antioxidant enzymes, osmolytes and also trigger their interactions with each other. However, research has been limited to only cross adaptations and signal transductions. Understanding the change and mechanism of H₂S and NO mediated cell signaling will broaden our knowledge on the various biochemical changes that occur in plant cells related to different stresses. A clear understanding of these molecules in various environmental stresses would help to confer biotechnological applications to protect plants against abiotic stresses and to improve crop productivity.