植物抗氧化动态平衡研究进展

植物在生长发育的过程中会产生代谢副产物活性氧,其含量在植物生长过程中起双重作用。适量的活性氧可提高植物对逆境胁迫的耐受性,但是过量的活性氧会诱发氧化猝发反应,严重影响植物的生长发育。因此,提高植物的抗氧化能力对于提高植物的抗逆能力来说显得尤为重要,该方面的研究也成为近年来逆境生物学的一大热点。植物体为了应对逆境环境造成的活性氧动态失衡,进化出了含酶和非酶组分的抗氧化系统。本文主要介绍了参与高等植物活性氧代谢的相关物质,对近年来国内外报道的代谢途径进行了综述,为提高植物的抗逆能力提供参考依据。     

Antioxidative response mechanisms in halophytes: Their role in stress defence

Normal growth and development of plants is greatly dependent on the capacity to overcome environmental stresses. Environmental stress conditions like high salinity, drought, high incident light and low or high temperature cause major crop losses worldwide. A common denominator in all these adverse conditions is the production of reactive oxygen species (ROS) within different cellular compartments of the plant cell. Plants have developed robust mechanisms including enzymatic or nonenzymatic scavenging pathways to counter the deleterious effects of ROS production. There are a number of general reviews on oxidative stress in plants and few on the role of ROS scavengers during stress conditions. Here we review the regulation of antioxidant enzymes during salt stress in halophytes, especially mangroves. We conclude that (i) antioxidant enzymes protect halophytes from deleterious ROS production during salt stress, and (ii) genetic information from mangroves and other halophytes would be helpful in defining the roles of individual isoforms. This information would be critical in using the appropriate genes for oxidative stress defence for genetic engineering of enhanced stress tolerance in crop systems.     

Biotechnological approach of improving plant salt tolerance using antioxidants as markers

Salt stress causes multifarious adverse effects in plants. Of them, production of reactive oxygen species (ROS) is a common phenomenon. These ROS are highly reactive because they can interact with a number of cellular molecules and metabolites thereby leading to a number of destructive processes causing cellular damage. Plants possess to a variable extent antioxidant metabolites, enzymes and non-enzymes, that have the ability to detoxify ROS. In the present review, the emphasis of discussion has been on understanding the role of different antioxidants in plants defense against oxidative stress caused by salt stress. The role of different antioxidants as potential selection criteria for improving plant salt tolerance has been critically discussed. With the advances in molecular biology and availability of advanced genetic tools considerable progress has been made in the past two decades in improving salt-induced oxidative stress tolerance in plants by developing transgenic lines with altered levels of antioxidants of different crops. The potential of this approach in counteracting stress-induced oxidative stress has been discussed at length in this review.     

Overexpression of dehydroascorbate reductase, but not monodehydroascorbate reductase, confers tolerance to aluminum stress in transgenic tobacco

Aluminum (Al) inhibits plant growth partly by causing oxidative damage that is promoted by reactive oxygen species and can be prevented by improving antioxidant capacity. Ascorbic acid (AsA), the most abundant antioxidant in plants, is regenerated by the action of monodehydroascorbate reductase (MDAR) and dehydroascorbate reductase (DHAR). We investigated the role of MDAR and DHAR in AsA regeneration during Al stress using transgenic tobacco (Nicotiana tabacum) plants overexpressing Arabidopsis cytosolic MDAR (MDAR-OX) or DHAR (DHAR-OX). DHAR-OX plants showed better root growth than wild-type (SR-1) plants after exposure to Al for 2 weeks, but MDAR-OX plants did not. There was no difference in Al distribution and accumulation in the root tips among SR-1, DHAR-OX, and MDAR-OX plants after Al treatment for 24 h. However, DHAR-OX plants showed lower hydrogen peroxide content, less lipid peroxidation and lower level of oxidative DNA damage than SR-1 plants, whereas MDAR-OX plants showed the same extent of damage as SR-1 plants. Compared with SR-1 plants, DHAR-OX plants consistently maintained a higher AsA level both with and without Al exposure, while MDAR-OX plants maintained a higher AsA level only without Al exposure. Also, DHAR-OX plants maintained higher APX activity under Al stress. The higher AsA level and APX activity in DHAR-OX plants contributed to their higher antioxidant capacity and higher tolerance to Al stress. These findings show that the overexpression of DHAR, but not of MDAR, confers Al tolerance, and that maintenance of a high AsA level is important to Al tolerance.     

The Involvement of Wheat F-Box Protein Gene TaFBA1 in the Oxidative Stress Tolerance of Plants

As one of the largest gene families, F-box domain proteins have been found to play important roles in abiotic stress responses via the ubiquitin pathway. TaFBA1 encodes a homologous F-box protein contained in E3 ubiquitin ligases. In our previous study, we found that the overexpression of TaFBA1 enhanced drought tolerance in transgenic plants. To investigate the mechanisms involved, in this study, we investigated the tolerance of the transgenic plants to oxidative stress. Methyl viologen was used to induce oxidative stress conditions. Real-time PCR and western blot analysis revealed that TaFBA1 expression was up-regulated by oxidative stress treatments. Under oxidative stress conditions, the transgenic tobacco plants showed a higher germination rate, higher root length and less growth inhibition than wild type (WT). The enhanced oxidative stress tolerance of the transgenic plants was also indicated by lower reactive oxygen species (ROS) accumulation, malondialdehyde (MDA) content and cell membrane damage under oxidative stress compared with WT. Higher activities of antioxidant enzymes, including superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and peroxidase (POD), were observed in the transgenic plants than those in WT, which may be related to the upregulated expression of some antioxidant genes via the overexpression of TaFBA1. In others, some stress responsive elements were found in the promoter region of TaFBA1, and TaFBA1 was located in the nucleus, cytoplasm and plasma membrane. These results suggest that TaFBA1 plays an important role in the oxidative stress tolerance of plants. This is important for understanding the functions of F-box proteins in plants’ tolerance to multiple stress conditions.     

The antioxidant system in Suaeda salsa under salt stress

ABSTRACTSuaeda salsaL. is a typical euhalophyte and is widely distributed throughout the world. Suaeda plants are important halophyte resources, and the physiological and biochemical characteristics of their various organsand their response to salt stress have been intensively studied. Leaf succulence, intracellular ion localization, increased osmotic regulation and enhanced antioxidant capacities are important responses for Suaeda plants to adapt to salt stress. Among these responses, scavenging of reactive oxygen species (ROS) is an important mechanism for plants to withstand oxidative stress and improve salt tolerance. The generation and scavenging pathways of ROS, as well as the expression of scavenging enzymes change under salt stress. This article reviews the antioxidant system constitute of S. salsa, and the mechanisms by which S. salsaantioxidant capacity is improved for salt tolerance. In addition, the differences between types of antioxidant mechanisms in S. salsaare reviewed, thereby revealing the adaptation mechanisms of Suaeda to different habitats. The review provides important clues for the comprehensive understanding of the salt tolerance mechanisms of halophytes.KEYWORDS: Suaeda salsa, halophyte, salt-tolerance mechanism, oxidative stress, antioxidant system     

The delayed leaf senescence mutants of Arabidopsis, ore1, ore3, and ore9 are tolerant to oxidative stress

Reactive oxygen species play a critical role in mediating the oxidative damage that causes senescence in a variety of aerobic organisms, from yeast to mammals. Genetic studies of these organisms have revealed that extended longevity is frequently associated with an increased resistance to stress. However, the relationship between life span and oxidative stress tolerance in plants is poorly understood. We have investigated the responses to oxidative stress in the delayed leaf senescence mutants of Arabidopsis thaliana, ore1, ore3, and ore9. The detached leaves of these mutants exhibit increased tolerance to various types of oxidative stress. The ore1, ore3, and ore9 mutants were also more tolerant to oxidative stress at the level of the whole plant, as determined by measuring physiological and molecular changes associated with oxidative stress. However, the activities of antioxidant enzymes were similar or lower in the mutants, as compared to wild type. These results suggest that the increased resistance to oxidative stress in the ore1, ore3, and ore9 mutants is not due to enhanced activities of these antioxidant enzymes. Taken together, our findings provide genetic evidence that oxidative stress tolerance is linked to control of leaf longevity in plants.     

Comparative Analysis of Physio-Biochemical Responses to Cold Stress in Tetraploid and Hexaploid Wheat

The cellular changes induced by cold stress (CS) include responses that lead to oxidative stress and limits plant growth, metabolism, and productivity. In this study, responses of physio-biochemical to CS phases were comparatively studied in three genotypes of bread and durum wheats differing in sensitivity, two of them (Norstar, bread wheat and Gerdish, durum wheat) were tolerant to CS and the other one, SRN (durum wheat) was sensitive to CS. 14-day-old seedlings were subjected to CS (12 and 24 h) with or without cold acclimation (CA) phase. During CS, the elevated levels of electrolyte leakage index, contents of hydrogen peroxide (H₂O₂), and malondialdehyde in Norstar and Gerdish were lower than that of SRN plants. Positive correlation and co-regulation of reactive oxygen species (ROS) scavenging systems, superoxide dismutase, catalase, ascorbate peroxidase, guaiacol peroxidase, and proline especially after CA phase suggested crucial role for holding back toxic ROS levels in CS phase. However, different activities of this system resulted in different intensities of oxidative stress in acclimated and non-acclimated plants. Our results showed that a CA phase induced oxidative stress tolerance by modulating antioxidative systems. These responses confirmed the existence of wide range of genetic capacity in durum wheat to increase cold tolerance particularly in Gerdish so that the sensitivity of SRN plants to CS was linearly correlated with the more decrease of antioxidant systems. These general responses may be a sign for associating other metabolites or enzymes activities to create partly tolerance against cold-induced oxidative stress. Eventually, assessing the dynamic of cell responses in short intervals after CS without CA phases profitably could be a novel path in plant stress response investigations in a short period of time.     

Antioxidant enzyme responses of plants to heavy metal stress

Heavy metal pollutions caused by natural processes or anthropological activities such as metal industries, mining, mineral fertilizers, pesticides and others pose serious environmental problems in present days. Evidently there is an urgent need of efficient remediation techniques that can tackle problems of such extent, especially in polluted soil and water resources. Phytoremediation is one such approach that devices effective and affordable ways of engaging suitable plants to cleanse the nature. Excessive accumulation of metal in plant tissues are known to cause oxidative stress. These, in turn differentially affect other plant processes that lead to loss of cellular homeostasis resulting in adverse affects on their growth and development apart from others. Plants have limited mechanisms of stress avoidance and require flexible means of adaptation to changing. A common feature to combat stress factors is synchronized function of antioxidant enzymes that helps alleviating cellular damage by limiting reactive oxygen species (ROS). Although, ROS are inevitable byproducts from essential aerobic metabolisms, these are needed under sub-lethal levels for normal plant growth. Understanding the interplay between oxidative stress in plants and role of antioxidant enzymes can result in developing plants that can overcome oxidative stress with the expression of antioxidant enzymes. These mechanisms have been proving to have immense potential for remediating these metals through the process of phytoremediation. The aim of this review is to assemble our current understandings of role of antioxidant enzymes of plants subjected to heavy metal stress.     

Plant responses to water stress: Role of reactive oxygen species

Terrestrial plants most often encounter drought stress because of erratic rainfall which has become compounded due to present climatic changes.Responses of plants to water stress may be assigned as either injurious change or tolerance index. One of the primary and cardinal changes in response to drought stress is the generation of reactive oxygen species (ROS), which is being considered as the cause of cellular damage. However, recently a signaling role of such ROS in triggering the ROS scavenging system that may confer protection or tolerance against stress is emerging. Such scavenging system consists of antioxidant enzymes like SOD, catalase and peroxidases, and antioxidant compounds like ascorbate, reduced glutathione; a balance between ROS generation and scavenging ultimately determines the oxidative load. As revealed in case of defence against pathogen, signaling via ROS is initiated by NADPH oxidase-catalyzed superoxide generation in the apoplastic space (cell wall) followed by conversion to hydrogen peroxide by the activity of cell wall-localized SOD. Wall peroxidase may also play role in ROS generation for signaling. Hydrogen peroxide may use Ca2+ and MAPK pathway as downstream signaling cascade. Plant hormones associated with stress responses like ABA and ethylene play their role possibly via a cross talk with ROS towards stress tolerance, thus projecting a dual role of ROS under drought stress.     

REACTIVE OXYGEN METABOLISM IN INTERTIDAL FUCUS SPP. (PHAEOPHYCEAE)

Our previous research suggests that interspecific variation in stress tolerance in intertidal Fucus spp. (Phaeophyceae) is partially mediated by differences in the production of, or ability to detoxify, reactive oxygen. Here we report on the content of antioxidants (ascorbate, glutathione, carotenoids, and tocopherols) and protective enzymes (catalase, superoxide dismutase, ascorbate peroxidase, and glutathione reductase) involved in reactive oxygen metabolism in three species of intertidal brown algae— Fucus spiralis L., F. evanescens C. Ag., and F. distichus L.—that differ in stress tolerance and position in the intertidal zone. Contents of the major antioxidants were similar in the three species and were not correlated with stress tolerance. The least stress tolerant species, F. distichus, had the lowest activity of reactive-oxygen-scavenging enzymes, although F. spiralis, the species with the highest stress tolerance, and F. evanescens contained similar activities of antioxidant enzymes on a fresh-weight basis. However, the activities of superoxide dismutase and ascorbate peroxidase in F. evanescens are lower than those of F. spiralis when expressed on the basis of chlorophyll. These data show that the ratio between reactive oxygen protection and production might be more important than the absolute content of antioxidants and protective enzymes. It also shows the importance of localization of detoxifying mechanisms and avoidance of oxidative stress.     

A Review on Selenium Function under Oxidative Stress in Plants Focusing on ROS Production and Detoxification

One of the main reasons of the annual reduction in plant production all around the world is the occurrence of abiotic stresses as a result of an unpredicted changes in environmental conditions. Abiotic stresses basically trigger numerous pathways related to oxygen free radicals’ generation resulting in a higher rate of reactive oxygen species (ROS) production. Accordingly, higher rate of oxygen free radicals than its steady state causes to oxidize various types of molecules and compartments within the plants’ cells and tissues. Oxidative stress is the result of high amount free radicals of oxygen interfering with different functions leading to undergo significant changes from molecular to phenotypic levels. In response to oxidative stress, plants deploy different enzymatic and non-enzymatic antioxidant mechanisms to detoxify extra free radicals and get back to a normal state. Applying some specific treatments have shown to significantly affect the antioxidant capacity and efficiency of the stressed cells and compartments. One of such reportedly effective treatments is the utilization of selenium (Se) element in stressed plants. Over the past years some different experiments evaluated the probable effect or efficiency of Se regarding its impact on plant under oxidative stress. Accordingly, based on the recent studies, Se has a significant role in plant responses to abiotic stresses probably due to its ability to improve the plants’ tolerance to oxidative stress. The significant influences of Se, and its related components such as nano-selenium, in plants under oxidative stress rooting from abiotic stresses, along with the new finding pertaining to its metabolism and translocation mechanisms inside the plant cells under oxidative stress condition are clearly explained in this review. However, there are still lack of a comprehensive explanation related to the precise mechanism of Se in plants under oxidative stress.

     

Effects of nitrogen on the activity of antioxidant enzymes and gene expression in leaves of Populus plants subjected to cadmium stress

The research aimed to verify the important physiological effect of nitrogen (N) on plants exposed to cadmium (Cd). The poplar plants were grown in a Hoagland nutrient solution and treated with extra N, Cd, and N + Cd. After treatment, plant growth and chlorophyll content were recorded. The oxidative stress, the activity of antioxidant enzymes, and the expression of related genes were also examined. The results indicated the plants treated with sole Cd presented obvious toxicity symptoms, i.e. growth inhibition, reactive oxygen species accumulation, and chlorophyll content decrement. However, when N was added to the plants under Cd stress, plant growth was enhanced, chlorophyll synthesis was promoted, and the oxidative stress was alleviated. Further, the expression of antioxidant enzymes genes was upregulated by N. The results indicated that N partially reversed the toxic effect of Cd on poplar plants, which can provide new methodology to enhance the phytoremediation technology for heavy metal pollution soil.     

Increased tolerance of rice to cold, drought and oxidative stresses mediated by the overexpression of a gene that encodes the zinc finger protein ZFP245

ZFP245 is a cold- and drought-responsive gene that encodes a zinc finger protein in rice. The ZFP245 protein localizes in the nucleus and exhibits trans-activation activity. Transgenic rice plants overexpressing ZFP245 were generated and found to display high tolerance to cold and drought stresses. The transgenic plants did not exhibit growth retardation, but showed growth sensitivity against exogenous abscisic acid, increased free proline levels and elevated expression of rice pyrroline-5-carboxylatesynthetase and proline transporter genes under stress conditions. Overproduction of ZFP245 enhanced the activities of reactive oxygen species-scavenging enzymes under stress conditions and increased the tolerance of rice seedlings to oxidative stress. Our data suggest that ZFP245 may contribute to the tolerance of rice plants to cold and drought stresses by regulating proline levels and reactive oxygen species-scavenging activities, and therefore may be useful for developing transgenic crops with enhanced tolerance to abiotic stress.     

Oxidant,antioxidant and physical exercise

Generation of reactive oxygen species (ROS) is a normal process in the life of aerobic organisms. Under physiological conditions, these deleterious species are mostly removed by the cellular antioxidant systems, which include antioxidant vitamins, protein and non-protein thiols, and antioxidant enzymes. Since the antioxidant reserve capacity in most tissues is rather marginal, strenuous physical exercise characterized by a remarkable increase in oxygen consumption with concomitant production of ROS presents a challenge to the antioxidant systems.An acute bout of exercise at sufficient intensity has been shown to stimulate activities of antioxidant enzymes. This could be considered as a defensive mechanism of the cell under oxidative stress. However, prolonged heavy exercise may cause a transient reduction of tissue vitamin E content and a change of glutathione redox status in various body tissues. Deficiency of antioxidant nutrients appears to hamper antioxidant systems and augment exercise-induced oxidative stress and tissue damage. Chronic exercise training seems to induce activities of antioxidant enzymes and perhaps stimulate GSH levels in body fluids. Recent research suggest that supplementation of certain antioxidant nutrients are necessary for physically active individuals.     

Colonization with arbuscular mycorrhizal fungus affects growth,drought tolerance and expression of stress-responsive genes in <Emphasis Type="Italic">Poncirus trifoliata</Emphasis>

The objective of this study was to investigate the effects of arbuscular mycorrhizal fungus (AMF) inoculation on growth and drought tolerance of Poncirus trifoliata seedlings. The seedlings were inoculated with or without Glomus mosseae before exposure to a short-term (3 days) water depletion, and relevant physiological and biochemical parameters (plant height, chlorophyll content, relative water content, activity of antioxidant enzymes) and expression patterns of several stress-responsive genes were examined. Inoculation with G. mosseae led to growth promotion of the seedlings, as revealed by larger plant height and higher relative water and chlorophyll contents. When subjected to drought treatment, the AMF-inoculated (AM) plants showed better tolerance than the nonmycorrhizal (NAM) plants. Under drought, the AM plants exhibited higher level of proline and activity of two antioxidant enzymes, superoxide dismutase (SOD) and peroxidase (POD). In addition, mRNA abundance of four genes involved in reactive oxygen species homeostasis and oxidative stress battling was higher in the AM plants when compared with the NAM plants. These results indicate that AMF inoculation stimulated growth and enhanced drought tolerance of the seedlings, which may be due to activation of an arsenal of physiological, biochemical and molecular alterations.     

Trichoderma harzianum enhances antioxidant defense of tomato seedlings and resistance to water deficit

Some plant-symbiotic strains of the genus Trichoderma colonize roots and induce profound changes in plant gene expression that lead to enhanced growth, especially under biotic and abiotic stresses. In this study, we tested the hypothesis that one of the protective mechanisms enhanced by T. harzianum T22 colonization is the antioxidant defense mechanism. Having established that strain T22 modulates the expression of the genes encoding antioxidant enzymes, the status of antioxidant defense of tomato seedlings in response to colonization by T22 and water deficit was investigated. Total ascorbate or glutathione levels were not affected by either stimuli, but under water deficit, antioxidant pools became more oxidized (lower ratios of reduced to oxidized forms), whereas colonized plants maintained redox state as high as or higher than unstressed and untreated plants. The enhanced redox state of colonized plants could be explained by their higher activity of ascorbate and glutathione-recycling enzymes, higher activity of superoxide dismutase, catalase, and ascorbate peroxidase, in both root and shoot throughout the experiment. Similar enzymes were induced in uncolonized plants in response to water-deficit stress but to a lower extent when compared with colonized plants. This orchestrated enhancement in activity of reactive oxygen species (ROS)-scavenging pathways in colonized plants in response to stress supports the hypothesis that enhanced resistance of colonized plants to water deficit is at least partly due to higher capacity to scavenge ROS and recycle oxidized ascorbate and glutathione, a mechanism that is expected to enhance tolerance to abiotic and biotic stresses.