油菜素内酯缓解番茄多环芳烃毒害及其解毒机理的研究
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摘要
多环芳烃(PAHs)是一种对环境中植物和动物持久危害的一类化合物。由于他们对人类的诱变致癌、致畸活性,多环芳烃已成为全球性关注的环境问题。由于它们广泛分布于土壤、空气和水中,从那里进入植物,从而对人类的健康构成威胁。这就是为什么研究对多环芳烃的解毒作用,从而确保食品安全的重要性。因此,研究植物对多环芳烃胁迫的抗性以及增强植物对其抗性及解毒机制对缓解植物多环芳烃毒害是非常重要的。油菜素内酯(BRs)是一种广谱性的植物特异性的内源甾体激素,可引起植物对各种非生物胁迫如高温、干旱、盐度、重金属的诱导抗性,并促进农药代谢。针对BRs的作用诱导抗性,本研究利用番茄对多环芳烃、镉(Cd)以及多氯联苯(PCBs)胁迫的反应,以及通过形态学、生理生化、分子和组织生物学的手段研究了24-表油菜素内酯(EBR)对不同生长期进和生育条件下的缓解效应。主要结果如下：
1.我们研究了不同浓度EBR (0.01、1、100nM)对不同浓度菲(PHE)(30、100、300μM)胁迫下番茄种子的发芽以及幼苗生长的影响。PHE胁迫延迟并降低了种子萌发率,处理10天后幼苗的高度和鲜重明显降低。EBR处理提高了种子萌发率以及幼苗鲜重,这种差异在PHE浓度为300μM表现最为显著。此外,与对照相比,EBR显著恢复了叶绿素荧光参数水平。在PHE胁迫下,各处理抗氧化酶表现不同,丙二醛(MDA)浓度明显增加。重要的是,EBR预处理增强了抗氧化酶活性并降低了MDA含量。综合考虑各研究参数,1nM EBR对缓解PHE胁迫的效果最好,其次为100nM和0.01nM。
2.研究表明,PHE和芘(PYR)处理的植株,随着PHE和PYR浓度的增加,处理植株的光合色素含量、光合速率、气孔导度、Fv/Fm、ΦPSⅡ、qP呈剂量依赖的方式下降,而NPQ是上升的,PHE比PYR处理产生了更为严重的胁迫。但是采用50nM和5nM的EBR处理叶片和根系显著提高了光合水平,缓解了胁迫。伴随PHE或PYR浓度的增加,SOD酶活性呈下降趋势,但GPOD、CAT、APX、GR及MDA的含量呈上升趋势。此外PHE或PYR处理降低了GSH和GSSG的含量。有趣的是PHE或PYR处理后再处理EBR可以提高根系抗氧化酶和非酶物质的含量。研究结果表明,番茄幼苗遭遇PHE或PYR侵害,EBR可通过增强植物的解毒活性起到抗污染胁迫的作用。
3.研究表明,以300μM的PHE处理21天时,通过气孔和非气孔因素,显著降低番茄幼苗的光合速率。叶绿素荧光参数的改变表明PHE处理发生了光抑制,进一步破坏光合作用机制。H2O2含量的积累及其在叶片中的定位表明,PHE处理产生了氧化胁迫。PHE处理诱导了与抗氧化相关的大多数酶活性。重要的是EBR+PHE处理比PHE单独处理显著增加了植株的生物量和光合速率,并且EBR+PHE处理比PHE单独处理更能够降低H202含量、增加抗氧化酶活性、促进解毒相关基因CYP90b3,GSH1和GST1的表达。有趣的是,PHE和PYR处理后,GSH含量和GSSG含量均呈降低趋势。相应GST含量的上升或许增加GSH的结合能力从而促进PHE的代谢。同时,EBR+PHE处理后,根系中较低的PHE含量表明PHE可能在根系中降解。结果表明,EBR通过植物解毒机制增强植物对PHE的抗性。
4.研究了PHE和EBR对番茄根系次生代谢和超微结构的影响。PHE处理增加了植物次生代谢相关酶GST、G6PDH、SKDH、PAL、CAD的活性,诱导相关基因的表达。EBR单独处理或者EBR和PHE共同处理诱导的次生代谢相关酶活性和基因表达均高于PHE单独处理。PHE处理诱导了酚类和类黄酮物质含量和相应酶含量的增加及DPPH的活性,而EBR处理则进一步增强了上述各项参数。PHE处理根尖细胞,细胞的超微结构发生改变,细胞核变形,线粒体数目减少,细胞壁变薄和较多的液泡,但是EBR处理后显著缓解了这些症状,细胞壁增厚,线粒体和内质网数目增加,液泡变小。结果表明EBR通过调节根尖次生代谢增强番茄根系耐受性。
5.重金属污染常伴随有机污染物,如PAHs,对这种协同污染的生理效应研究较少。但对混合污染的胁迫反应及抗性机理对植物的生长发育和抗性增强是非常重要的。因此,我们在温室内研究了100μM PHE和100μM Cd(?)昆合肋、迫效应以及EBR的缓解效应。处理40d后,PHE处理、Cd处理以及协同污染显著降低了同化物合成、光合作用以及叶绿素含量。让人出乎意料的是,PHE、Cd协同污染并没有降低生长量,相反却稍微提高了干物质量,光合速率和叶绿素含量。单一或复合污染提高了H202和MDA的含量、抗氧化物活性及相关基因表达,表明以上污染导致了氧化胁迫。另外PHE处理和Cd处理诱导了番茄叶片中次生代谢产物,解毒相关酶的活性及谷胱甘肽含量的提高。有趣的是,对胁迫植株外源喷施EBR后,可有效提高其生物量、光合、叶绿素,但降低了H202和MDA含量,同时伴随有抗氧化酶活性的提高。另外EBR激活了与次生代谢和解毒相关酶的活性,从而提高了其抗污染的能力。我们的结果表明,外源喷施EBR通过有效提高抗氧化能力、解毒能力和次生代谢从而提高混合污染胁迫抗性。
6. PAHs能够影响多种种植的蔬菜植物,PAHs对蔬菜植物各种生理参数的影响及其植物对PAHs胁迫的响应是胁迫生物学重要的研究内容。PHE处理5种蔬菜植物14天叶龄的叶片,叶片的生长量、光合作用和叶绿素含量以剂量依赖的方式均呈现下降趋势,MDA和H202以剂量依赖的方式呈上升趋势,抗氧化酶活性上升,表明PHE产生了氧化胁迫。但是,在一定范围,不同蔬菜种类的生长量和抗氧化防御有所差异,从而导致植物产生不同的耐受性和植物毒性。总之,5种蔬菜植物遭遇PHE毒性顺序为小白菜、黄瓜、生菜、番茄和菜苔。
7. PCBs是空气中常见的永久性有机污染物。空气中的PCBs的植物修复从而减轻对人体的伤害是一个新概念。然而空气中的PCBs对植物生长、光合和抗氧化系统研究较少。同时BRs已被作为一种潜在的植物调控相关的生理激素。因此,我们研究了PCBs和EBR对同化产物积累、光合机理、抗氧化酶活性的影响。不同浓度(0.4,2.0,10μg/L)的喷施显著降低了植株干重、光合速率、叶绿素含量。PCBs诱导的光合抑制与气孔、非气孔因素均相关。同时,不同水平的PCBs处理Fv/Fm、 ΦPSⅡ、 qP均明显降低,表明PCBs可诱导光抑制。H202和MDA的累计表明幼苗在PCBs的胁迫下遭受了氧化胁迫,进而产生了过量的ROS导致了抗氧化酶活性受抑制。相反,喷施100nmEBR提高了同化物质积累、光合速率、叶绿素含量、Fv/Fm、ΦPSⅡ、qP、进而降低了光抑制,EBR通过激活抗氧化活性减轻了ROS积累和MDA毒害。我们的结果表明,EBR具有对PCBs胁迫的保护作用,它通过提高植物抗性,进而加强了植物修复。
Polycyclic aromatic hydrocarbons (PAHs) are environmentally persistent organic micro pollutants. PAHs are of global environmental concerns because of their mutagenic, carcinogenic and teratogenic activities in humans. They are ubiquitous commonly found in soil, air and water, from where they enter plants, contaminate food chain and thus pose threat to human health. That is why, in planta detoxification of PAHs is very important to ensure food safety. Again, plants are important in removing PAHs, yet, understanding stress responses as well as strengthening plant tolerance and detoxification systems are important for plant based remediation of PAHs. Plant tolerance to PAHs is rate or capacity limiting as this is dependent on antioxidant and detoxification potential of plant. Hence, strengthening antioxidant and detoxification systems by hormonal supplementation may improve plant tolerance and degradation potential of xenobiotics like PAHs. Brassinosteroids (BRs), a class of plant-specific essential steroidal hormones are well known to induce plant tolerance against various abiotic stresses like high temperature, drought, salinity, heavy metal stress and also promote pesticide metabolism. Keeping in view the anti-stress properties of BRs, present study was carried out to understand the stress responses of tomato plant to PAHs (phenanthrene and pyrene), cadmium (Cd) and polychlorinated biphenyls (PCBs) as well as role of24-epibrassinolide (EBR) in stress amelioration through a series of experiments in different stages and conditions using morphological, physiological, biochemical, molecular and ultrastructural approaches. The salient findings are as follows:
(1) The present experiment investigated the effects of seed treatment with various concentrations (0.01,1.0,100nM) of EBR on seed germination and early seedling growth in tomato under graded levels (30,100,300μM) of phenanthrene (PHE). Delayed and decreased seed germination; reduced length and fresh weight (FW) of shoot and root were observed following10days of PHE exposure in a dose dependent manner. However, seed treatment with EBR improved seed germination and increased length and FW of shoot and root. In addition, EBR remarkably restored the studied chlorophyll fluorescence parameters towards control levels. Different responses in antioxidant enzymes were observed following exposure to PHE, while malondialdehyde (MDA) was increased in a concentration dependent manner. Importantly, EBR pretreatment further increased activities of antioxidant enzymes but decreased the MDA content both in shoot and root of young tomato seedlings. Considering all the studied parameters, seed treatment with1.0nM EBR was most effective followed by100nM and0.01nM for improvement of germination and seedling growth under PHE stress in tomato.
(2) A concentration-dependent decrease in growth, photosynthetic pigment contents, net photosynthetic rate (Pn), stomatal conductance (Gs), maximal quantum yield of PSII (Fv/Fm), effective quantum yield of PSⅡ (ΦPSⅡ) and photochemical quenching coefficient (qP) has been observed following phenanthrene (PHE) and pyrene (PYR) exposure (0,10,30,100and300μM). By contrast, non-photochemical quenching coefficient (NPQ) was increased. PHE was found to induce higher stress than PYR. However, foliar or root application of EBR (50nM and5nM, respectively) alleviated all those depressions with a sharp improvement in the activity of photosynthetic machinery. Superoxide dismutase (SOD) activity was gradually declined with increased concentration of PHE or PYR. However, the activities of guaiacol peroxidase (GPOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR) as well as content of MDA were increased in a dose-dependent manner. In addition, both reduced glutathione (GSH) and oxidized glutathione (GSSG) were induced by PHE or PYR. Interestingly, EBR application in either form further increased enzymatic and non enzymatic antioxidants in tomato roots treated with PHE or PYR. Our results suggest that EBR has an anti-stress effect on tomato seedlings contaminated with PHE or PYR and this effect is mainly attributed by increased detoxification activity.
(3) Exposure to PHE (300μM) for21d significantly decreased Pn which was attributed by both stomatal and nonstomatal factors. Changes in chlorophyll fluorescence parameters showed that PHE induced photoinhibition and subsequent damage to the photosynthetic machinery. Increased H2O2contents and localized H2O2accumulation in leaves indicated PHE-induced oxidative damage. All most all of the studied antioxidant enzymes were induced by PHE. Importantly, foliar application of EBR (0.1μM) significantly increased Pn and plant biomass over PHE alone. Furthermore, EBR+PHE showed remarkably decreased H2O2contents accompanied with increased antioxidant enzymes activities over PHE alone. Enhanced expression of detoxification related genes CYP90b3, GSH1and GST1suggested a strong coordinated detoxification under EBR+PHE treatment. Increased GSH and decreased GSSG contents resulted high GSH/GSSG in EBL+PHE treatment. In agreement with gene expression, a concurrent increment of glutathione S-transferase (GST) activity in EBL+PHE treatment over PHE alone was observed. Increased GST activity might promote conjugation of PHE with GSH and thereby improving metabolism of PHE. Meanwhile, remarkably lower PHE residue in root under EBR+PHE treatment suggested possible PHE degradation in root. Our results indicate that EBR application could be an effective tool to enhance plant tolerance against PHE stress via in planta detoxification.
(4) This experiment was carried out to investigate the effects of PHE and EBR on secondary metabolism and ultra structure in tomato root. Exposure to PHE increased the activity of secondary metabolism related enzymes activities viz. GST, glucose-6-phosphate dehydrogenase (G6PDH), shikimate dehydrogenase (SKDH), phenylalanine ammonialyase (PAL) and cinnamyl alcohol dehydrogenase (CAD). The expressions of related genes were also induced by PHE. Meanwhile, EBR alone or in combination with PHE increased the enzymes activity and gene expression significantly over control and PHE alone, respectively. Consistent with enzymes activities contents of phenols, flavonoids and activity of DPPH were induced by PHE, while EBR application further increased all those studied parameters. Obvious ultrastructural alterations characterized by irregular shaped nucleus, less mitochondria, thin cell wall and more vacuolation were observed in PHE-treated root tip cells. However, EBR application saved root cell from severe PHE-induced damage which was clearly observed by relatively thick cell wall, more mitochondria, endoplasmic reticulum and small size vacuole in EBR+PHE-treated root tip cells. These observations suggest that EBR regulates secondary metabolism in tomato which might enhance tolerance to PHE.
(5) Environmental pollution by heavy metal is often occurred with organic contaminants such as PAHs. However, plant stress physiology under co-contamination is poorly understood area. But knowledge on stress responses and stress tolerance is very important to develop and strengthen plant based remediation of mixed pollution as well as for food safety. Therefore, a greenhouse experiment was conducted to understand the stress responses of PHE and/or Cd to tomato plant and role of EBR in stress amelioration. PHE or Cd alone or in combination significantly reduced the biomass, Pn and photosynthetic pigment contents in tomato plant after40d of respective treatments. Surprisingly, combination of PHE and Cd did not reduce the growth synergistically, rather slightly improved the dry weight, Pn and leaf pigments. Single or dual pollution increased H2O2, MDA content, antioxidant enzymes activity and related genes expression which indicate an occurrence of oxidative stress. In addition, PHE and/or Cd induced the secondary metabolism and detoxification related enzymes activity as well as glutathione contents in tomato leaves. The foliar application of EBR on stressed plants, increased biomass, Pn, leaf pigments, but decreased H2O2and MDA contents by the action of enhanced antioxidant enzymes activity. EBR stimulates the secondary metabolism and xenobiotics detoxification related enzymes activity as well as gene expression towards enhanced tolerance to pollutant stress. Additionally, PHE and/or Cd residues were significantly decreased both in leaves and roots after application of EBR that indicate a possible promotion in degradation and detoxification of pollutants. Our results suggest that EBR could alleviate PHE-Cd co-contamination-induced stress by enhancing photosynthesis, antioxidant defense, secondary metabolism and pollutant detoxification capacity.
(6) Cultivated vegetable species are continuously exposed to atmospheric PAHs; yet, information on their effects on different physiological parameters and related stress responses is an important part of stress biology. Two weeks foliar exposure of PHE resulted more or less dose dependent decrease in growth, photosynthesis and chlorophyll contents in five studied vegetables. With few exceptions, activities of antioxidant enzymes were upregulated following PHE exposure. Dose dependent increase in MDA contents together with H2O2accumulation suggested an occurrence of oxidative stress by PHE. However, to some extent, growth and antioxidant defense responses differ from species to species. Difference in defense capacity might result different tolerance and phytotoxicity among the studied vegetables. Taken together, phytotoxicity of PHE to five vegetables could be sequenced in the following order: pakchoi>cucumber>lettuce>tomato>chinese flowering cabbage.
(7) PCBs are persistent organic pollutants often found in the atmosphere. Phytoremediation of airborne PCBs is an emerging new concept to minimize potential human exposure. However, effects of atmospheric PCBs on plant growth, photosynthesis and antioxidant defence system are poorly understood area. Meanwhile, BRs have been proposed as potential phytohormone for "Assisted phytoremediation by plant growth regulators" program. Hence, we studied the effects of PCBs and EBR on biomass accumulation, photo synthetic machinery and antioxidant system in tomato plants. PCBs (0.4,2.0and10μg/1) mist spray significantly decreased dry weight, photosynthesis, chlorophyll contents in a dose dependent manner. Both stomatal and non stomatal factors are involved in PCBs-induced photosynthetic inhibition. Meanwhile, Fv/Fm, ΦPSⅡ and qP were increasingly decreased by various levels of PCBs, suggested an induction of photoinhibition. Increased accumulation of H2O2accompanied with high lipid peroxidation suggested that seedlings faced oxidative stress upon PCBs exposure. Accordingly, antioxidant enzyme activity was inhibited due to inactivation of enzymes by over production of reactive oxygen species (ROS). In contrast, EBR (100nM) application increased biomass, photosynthetic capacity, chlorophyll contents and alleviated photoinhibition by enhancing Fv/Fm, ΦPSⅡ and qP. EBR significantly decreased harmful ROS accumulation and lipid peroxidation through upregulating antioxidant enzymes activity. Our results suggest a protective role of EBR against PCBs stress which may strengthen phytoremediation approaches by enhancing plant tolerance.
1.我们研究了不同浓度EBR (0.01、1、100nM)对不同浓度菲(PHE)(30、100、300μM)胁迫下番茄种子的发芽以及幼苗生长的影响。PHE胁迫延迟并降低了种子萌发率,处理10天后幼苗的高度和鲜重明显降低。EBR处理提高了种子萌发率以及幼苗鲜重,这种差异在PHE浓度为300μM表现最为显著。此外,与对照相比,EBR显著恢复了叶绿素荧光参数水平。在PHE胁迫下,各处理抗氧化酶表现不同,丙二醛(MDA)浓度明显增加。重要的是,EBR预处理增强了抗氧化酶活性并降低了MDA含量。综合考虑各研究参数,1nM EBR对缓解PHE胁迫的效果最好,其次为100nM和0.01nM。
2.研究表明,PHE和芘(PYR)处理的植株,随着PHE和PYR浓度的增加,处理植株的光合色素含量、光合速率、气孔导度、Fv/Fm、ΦPSⅡ、qP呈剂量依赖的方式下降,而NPQ是上升的,PHE比PYR处理产生了更为严重的胁迫。但是采用50nM和5nM的EBR处理叶片和根系显著提高了光合水平,缓解了胁迫。伴随PHE或PYR浓度的增加,SOD酶活性呈下降趋势,但GPOD、CAT、APX、GR及MDA的含量呈上升趋势。此外PHE或PYR处理降低了GSH和GSSG的含量。有趣的是PHE或PYR处理后再处理EBR可以提高根系抗氧化酶和非酶物质的含量。研究结果表明,番茄幼苗遭遇PHE或PYR侵害,EBR可通过增强植物的解毒活性起到抗污染胁迫的作用。
3.研究表明,以300μM的PHE处理21天时,通过气孔和非气孔因素,显著降低番茄幼苗的光合速率。叶绿素荧光参数的改变表明PHE处理发生了光抑制,进一步破坏光合作用机制。H2O2含量的积累及其在叶片中的定位表明,PHE处理产生了氧化胁迫。PHE处理诱导了与抗氧化相关的大多数酶活性。重要的是EBR+PHE处理比PHE单独处理显著增加了植株的生物量和光合速率,并且EBR+PHE处理比PHE单独处理更能够降低H202含量、增加抗氧化酶活性、促进解毒相关基因CYP90b3,GSH1和GST1的表达。有趣的是,PHE和PYR处理后,GSH含量和GSSG含量均呈降低趋势。相应GST含量的上升或许增加GSH的结合能力从而促进PHE的代谢。同时,EBR+PHE处理后,根系中较低的PHE含量表明PHE可能在根系中降解。结果表明,EBR通过植物解毒机制增强植物对PHE的抗性。
4.研究了PHE和EBR对番茄根系次生代谢和超微结构的影响。PHE处理增加了植物次生代谢相关酶GST、G6PDH、SKDH、PAL、CAD的活性,诱导相关基因的表达。EBR单独处理或者EBR和PHE共同处理诱导的次生代谢相关酶活性和基因表达均高于PHE单独处理。PHE处理诱导了酚类和类黄酮物质含量和相应酶含量的增加及DPPH的活性,而EBR处理则进一步增强了上述各项参数。PHE处理根尖细胞,细胞的超微结构发生改变,细胞核变形,线粒体数目减少,细胞壁变薄和较多的液泡,但是EBR处理后显著缓解了这些症状,细胞壁增厚,线粒体和内质网数目增加,液泡变小。结果表明EBR通过调节根尖次生代谢增强番茄根系耐受性。
5.重金属污染常伴随有机污染物,如PAHs,对这种协同污染的生理效应研究较少。但对混合污染的胁迫反应及抗性机理对植物的生长发育和抗性增强是非常重要的。因此,我们在温室内研究了100μM PHE和100μM Cd(?)昆合肋、迫效应以及EBR的缓解效应。处理40d后,PHE处理、Cd处理以及协同污染显著降低了同化物合成、光合作用以及叶绿素含量。让人出乎意料的是,PHE、Cd协同污染并没有降低生长量,相反却稍微提高了干物质量,光合速率和叶绿素含量。单一或复合污染提高了H202和MDA的含量、抗氧化物活性及相关基因表达,表明以上污染导致了氧化胁迫。另外PHE处理和Cd处理诱导了番茄叶片中次生代谢产物,解毒相关酶的活性及谷胱甘肽含量的提高。有趣的是,对胁迫植株外源喷施EBR后,可有效提高其生物量、光合、叶绿素,但降低了H202和MDA含量,同时伴随有抗氧化酶活性的提高。另外EBR激活了与次生代谢和解毒相关酶的活性,从而提高了其抗污染的能力。我们的结果表明,外源喷施EBR通过有效提高抗氧化能力、解毒能力和次生代谢从而提高混合污染胁迫抗性。
6. PAHs能够影响多种种植的蔬菜植物,PAHs对蔬菜植物各种生理参数的影响及其植物对PAHs胁迫的响应是胁迫生物学重要的研究内容。PHE处理5种蔬菜植物14天叶龄的叶片,叶片的生长量、光合作用和叶绿素含量以剂量依赖的方式均呈现下降趋势,MDA和H202以剂量依赖的方式呈上升趋势,抗氧化酶活性上升,表明PHE产生了氧化胁迫。但是,在一定范围,不同蔬菜种类的生长量和抗氧化防御有所差异,从而导致植物产生不同的耐受性和植物毒性。总之,5种蔬菜植物遭遇PHE毒性顺序为小白菜、黄瓜、生菜、番茄和菜苔。
7. PCBs是空气中常见的永久性有机污染物。空气中的PCBs的植物修复从而减轻对人体的伤害是一个新概念。然而空气中的PCBs对植物生长、光合和抗氧化系统研究较少。同时BRs已被作为一种潜在的植物调控相关的生理激素。因此,我们研究了PCBs和EBR对同化产物积累、光合机理、抗氧化酶活性的影响。不同浓度(0.4,2.0,10μg/L)的喷施显著降低了植株干重、光合速率、叶绿素含量。PCBs诱导的光合抑制与气孔、非气孔因素均相关。同时,不同水平的PCBs处理Fv/Fm、 ΦPSⅡ、 qP均明显降低,表明PCBs可诱导光抑制。H202和MDA的累计表明幼苗在PCBs的胁迫下遭受了氧化胁迫,进而产生了过量的ROS导致了抗氧化酶活性受抑制。相反,喷施100nmEBR提高了同化物质积累、光合速率、叶绿素含量、Fv/Fm、ΦPSⅡ、qP、进而降低了光抑制,EBR通过激活抗氧化活性减轻了ROS积累和MDA毒害。我们的结果表明,EBR具有对PCBs胁迫的保护作用,它通过提高植物抗性,进而加强了植物修复。
Polycyclic aromatic hydrocarbons (PAHs) are environmentally persistent organic micro pollutants. PAHs are of global environmental concerns because of their mutagenic, carcinogenic and teratogenic activities in humans. They are ubiquitous commonly found in soil, air and water, from where they enter plants, contaminate food chain and thus pose threat to human health. That is why, in planta detoxification of PAHs is very important to ensure food safety. Again, plants are important in removing PAHs, yet, understanding stress responses as well as strengthening plant tolerance and detoxification systems are important for plant based remediation of PAHs. Plant tolerance to PAHs is rate or capacity limiting as this is dependent on antioxidant and detoxification potential of plant. Hence, strengthening antioxidant and detoxification systems by hormonal supplementation may improve plant tolerance and degradation potential of xenobiotics like PAHs. Brassinosteroids (BRs), a class of plant-specific essential steroidal hormones are well known to induce plant tolerance against various abiotic stresses like high temperature, drought, salinity, heavy metal stress and also promote pesticide metabolism. Keeping in view the anti-stress properties of BRs, present study was carried out to understand the stress responses of tomato plant to PAHs (phenanthrene and pyrene), cadmium (Cd) and polychlorinated biphenyls (PCBs) as well as role of24-epibrassinolide (EBR) in stress amelioration through a series of experiments in different stages and conditions using morphological, physiological, biochemical, molecular and ultrastructural approaches. The salient findings are as follows:
(1) The present experiment investigated the effects of seed treatment with various concentrations (0.01,1.0,100nM) of EBR on seed germination and early seedling growth in tomato under graded levels (30,100,300μM) of phenanthrene (PHE). Delayed and decreased seed germination; reduced length and fresh weight (FW) of shoot and root were observed following10days of PHE exposure in a dose dependent manner. However, seed treatment with EBR improved seed germination and increased length and FW of shoot and root. In addition, EBR remarkably restored the studied chlorophyll fluorescence parameters towards control levels. Different responses in antioxidant enzymes were observed following exposure to PHE, while malondialdehyde (MDA) was increased in a concentration dependent manner. Importantly, EBR pretreatment further increased activities of antioxidant enzymes but decreased the MDA content both in shoot and root of young tomato seedlings. Considering all the studied parameters, seed treatment with1.0nM EBR was most effective followed by100nM and0.01nM for improvement of germination and seedling growth under PHE stress in tomato.
(2) A concentration-dependent decrease in growth, photosynthetic pigment contents, net photosynthetic rate (Pn), stomatal conductance (Gs), maximal quantum yield of PSII (Fv/Fm), effective quantum yield of PSⅡ (ΦPSⅡ) and photochemical quenching coefficient (qP) has been observed following phenanthrene (PHE) and pyrene (PYR) exposure (0,10,30,100and300μM). By contrast, non-photochemical quenching coefficient (NPQ) was increased. PHE was found to induce higher stress than PYR. However, foliar or root application of EBR (50nM and5nM, respectively) alleviated all those depressions with a sharp improvement in the activity of photosynthetic machinery. Superoxide dismutase (SOD) activity was gradually declined with increased concentration of PHE or PYR. However, the activities of guaiacol peroxidase (GPOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR) as well as content of MDA were increased in a dose-dependent manner. In addition, both reduced glutathione (GSH) and oxidized glutathione (GSSG) were induced by PHE or PYR. Interestingly, EBR application in either form further increased enzymatic and non enzymatic antioxidants in tomato roots treated with PHE or PYR. Our results suggest that EBR has an anti-stress effect on tomato seedlings contaminated with PHE or PYR and this effect is mainly attributed by increased detoxification activity.
(3) Exposure to PHE (300μM) for21d significantly decreased Pn which was attributed by both stomatal and nonstomatal factors. Changes in chlorophyll fluorescence parameters showed that PHE induced photoinhibition and subsequent damage to the photosynthetic machinery. Increased H2O2contents and localized H2O2accumulation in leaves indicated PHE-induced oxidative damage. All most all of the studied antioxidant enzymes were induced by PHE. Importantly, foliar application of EBR (0.1μM) significantly increased Pn and plant biomass over PHE alone. Furthermore, EBR+PHE showed remarkably decreased H2O2contents accompanied with increased antioxidant enzymes activities over PHE alone. Enhanced expression of detoxification related genes CYP90b3, GSH1and GST1suggested a strong coordinated detoxification under EBR+PHE treatment. Increased GSH and decreased GSSG contents resulted high GSH/GSSG in EBL+PHE treatment. In agreement with gene expression, a concurrent increment of glutathione S-transferase (GST) activity in EBL+PHE treatment over PHE alone was observed. Increased GST activity might promote conjugation of PHE with GSH and thereby improving metabolism of PHE. Meanwhile, remarkably lower PHE residue in root under EBR+PHE treatment suggested possible PHE degradation in root. Our results indicate that EBR application could be an effective tool to enhance plant tolerance against PHE stress via in planta detoxification.
(4) This experiment was carried out to investigate the effects of PHE and EBR on secondary metabolism and ultra structure in tomato root. Exposure to PHE increased the activity of secondary metabolism related enzymes activities viz. GST, glucose-6-phosphate dehydrogenase (G6PDH), shikimate dehydrogenase (SKDH), phenylalanine ammonialyase (PAL) and cinnamyl alcohol dehydrogenase (CAD). The expressions of related genes were also induced by PHE. Meanwhile, EBR alone or in combination with PHE increased the enzymes activity and gene expression significantly over control and PHE alone, respectively. Consistent with enzymes activities contents of phenols, flavonoids and activity of DPPH were induced by PHE, while EBR application further increased all those studied parameters. Obvious ultrastructural alterations characterized by irregular shaped nucleus, less mitochondria, thin cell wall and more vacuolation were observed in PHE-treated root tip cells. However, EBR application saved root cell from severe PHE-induced damage which was clearly observed by relatively thick cell wall, more mitochondria, endoplasmic reticulum and small size vacuole in EBR+PHE-treated root tip cells. These observations suggest that EBR regulates secondary metabolism in tomato which might enhance tolerance to PHE.
(5) Environmental pollution by heavy metal is often occurred with organic contaminants such as PAHs. However, plant stress physiology under co-contamination is poorly understood area. But knowledge on stress responses and stress tolerance is very important to develop and strengthen plant based remediation of mixed pollution as well as for food safety. Therefore, a greenhouse experiment was conducted to understand the stress responses of PHE and/or Cd to tomato plant and role of EBR in stress amelioration. PHE or Cd alone or in combination significantly reduced the biomass, Pn and photosynthetic pigment contents in tomato plant after40d of respective treatments. Surprisingly, combination of PHE and Cd did not reduce the growth synergistically, rather slightly improved the dry weight, Pn and leaf pigments. Single or dual pollution increased H2O2, MDA content, antioxidant enzymes activity and related genes expression which indicate an occurrence of oxidative stress. In addition, PHE and/or Cd induced the secondary metabolism and detoxification related enzymes activity as well as glutathione contents in tomato leaves. The foliar application of EBR on stressed plants, increased biomass, Pn, leaf pigments, but decreased H2O2and MDA contents by the action of enhanced antioxidant enzymes activity. EBR stimulates the secondary metabolism and xenobiotics detoxification related enzymes activity as well as gene expression towards enhanced tolerance to pollutant stress. Additionally, PHE and/or Cd residues were significantly decreased both in leaves and roots after application of EBR that indicate a possible promotion in degradation and detoxification of pollutants. Our results suggest that EBR could alleviate PHE-Cd co-contamination-induced stress by enhancing photosynthesis, antioxidant defense, secondary metabolism and pollutant detoxification capacity.
(6) Cultivated vegetable species are continuously exposed to atmospheric PAHs; yet, information on their effects on different physiological parameters and related stress responses is an important part of stress biology. Two weeks foliar exposure of PHE resulted more or less dose dependent decrease in growth, photosynthesis and chlorophyll contents in five studied vegetables. With few exceptions, activities of antioxidant enzymes were upregulated following PHE exposure. Dose dependent increase in MDA contents together with H2O2accumulation suggested an occurrence of oxidative stress by PHE. However, to some extent, growth and antioxidant defense responses differ from species to species. Difference in defense capacity might result different tolerance and phytotoxicity among the studied vegetables. Taken together, phytotoxicity of PHE to five vegetables could be sequenced in the following order: pakchoi>cucumber>lettuce>tomato>chinese flowering cabbage.
(7) PCBs are persistent organic pollutants often found in the atmosphere. Phytoremediation of airborne PCBs is an emerging new concept to minimize potential human exposure. However, effects of atmospheric PCBs on plant growth, photosynthesis and antioxidant defence system are poorly understood area. Meanwhile, BRs have been proposed as potential phytohormone for "Assisted phytoremediation by plant growth regulators" program. Hence, we studied the effects of PCBs and EBR on biomass accumulation, photo synthetic machinery and antioxidant system in tomato plants. PCBs (0.4,2.0and10μg/1) mist spray significantly decreased dry weight, photosynthesis, chlorophyll contents in a dose dependent manner. Both stomatal and non stomatal factors are involved in PCBs-induced photosynthetic inhibition. Meanwhile, Fv/Fm, ΦPSⅡ and qP were increasingly decreased by various levels of PCBs, suggested an induction of photoinhibition. Increased accumulation of H2O2accompanied with high lipid peroxidation suggested that seedlings faced oxidative stress upon PCBs exposure. Accordingly, antioxidant enzyme activity was inhibited due to inactivation of enzymes by over production of reactive oxygen species (ROS). In contrast, EBR (100nM) application increased biomass, photosynthetic capacity, chlorophyll contents and alleviated photoinhibition by enhancing Fv/Fm, ΦPSⅡ and qP. EBR significantly decreased harmful ROS accumulation and lipid peroxidation through upregulating antioxidant enzymes activity. Our results suggest a protective role of EBR against PCBs stress which may strengthen phytoremediation approaches by enhancing plant tolerance.
引文
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