内生细菌在重金属植物修复中的作用机理及应用研究
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摘要
随着工业生产的发展，有毒重金属污染越来越受到关注。重金属能在生物体内累积，最终通过食物链危害人类的健康。传统的重金属污染土壤修复技术通常都耗费较高而且对环境有较大伤害。因此，急需一种行之有效且经济的去除环境中重金属的方法。原位植物修复技术因其成本低、环境友好且行之有效而受到国内外研究者的关注，被认为是一种极具前景的污染环境修复方法。然而，重金属的植物修复的实际应用还有不少问题亟待解决，譬如重金属植物毒性、修复周期长且污染物提取量有限等。现有的植物修复强化措施（譬如转基因和螯合剂等）虽然有效但费用昂贵且对环境有潜在风险。因此，寻求一条经济且有效的微生物强化途径提高植物修复效率将意义重大。
本文自镉超累积植物龙葵中分离得到多株内生菌，通过多种性能指标筛选菌株Serratia nematodiphila LRE07作为模式菌株，用以研究利用植物-内生菌共生系统强化植物修复的可行性。将菌株S. nematodiphila LRE07接种到龙葵种子后，通过营养液培养方法深入研究了系列镉处理浓度下龙葵植株生长、生理参数、光合成色素和镉吸收累积特性，探讨了内生菌接种对植物抗压性能的影响。在此基础上，研究了镉胁迫下植物对营养元素的摄取、脂质过氧化水平和活性氧代谢的变化，以揭示内生菌在降低重金属植物毒性和促进植物生长中的可能作用机制。另外，还研究了菌株S. nematodiphila LRE07的稳定性和其在非宿主植物体内的作用效果。主要的研究内容和结果如下：
①龙葵内生菌的分离与筛选
用于内生菌分离的龙葵植株采自株洲冶炼厂（27°52′N,113°05′E）污水排水渠堤，自该植物不同组织体内分离得到97株内生菌，其中大部分为细菌。通过最小抑制浓度测定发现所有内生菌对重金属均有不同程度抗性，这表明超累积植物是筛选重金属抗性菌株的天然资源。自龙葵根部得到一株产深红色色素的细菌，命名为LRE07。通过16S rDNA分析鉴定该菌株为沙雷氏菌株（Serratianematodiphila）。S. nematodiphila LRE07表现出对多种重金属均有较高抗性，尤其是对铜、镉和铬。同时，对该菌株进行植物促生相关的生物学性能测定，结果发现S. nematodiphila LRE07具有产吲哚乙酸、铁载体和增溶矿质磷酸盐的能力。上述性能表明该菌株具有重金属抗性和植物促生潜力，因此确定S. nematodiphilaLRE07为下一步实验菌株。
②内生菌接种对龙葵植株生长和镉吸收的影响
通过营养液培养方法，研究龙葵植株对镉胁迫的响应和内生菌接种对植株抗压性能的影响。接种21天和70天后取植物样测定菌株接种效率。结果表明内生菌S. nematodiphila LRE07成功接种入植株体内，自宿主植物根部和地上部分组织内重分离得到的菌落数表现出依赖时间增长的趋势，镉胁迫未对内生菌生长造成抑制作用。上述结果表明S. nematodiphila LRE07能与宿主植物和谐共生。镉浓度的递增对龙葵生长造成了不同程度抑制作用，表现在植株的分蘖数、鲜重和株高。随着重金属接触时间延长，0μM和10μM浓度下接种植物和未接种植物的分蘖数比率均有增加，但高浓度（50和100μM）下分蘖数比率很快达到恒定值甚至在实验后期减少。营养液中重金属浓度越高，植物的分蘖数比率越低。在同一镉胁迫浓度下，接种内生菌的植株分蘖数比率要高于未接种植株（P <0.05,MANOVAs），即接种植株的叶片数更多。内生菌接种对植株生长的有利影响在植株鲜重、株高和根重上也有体现。内生菌的积极意义在低镉胁迫（10μM）下尤为显著，在该浓度下，接种植物生长未见显著抑制，其生物量比未接种植株提高了58%。
内生菌的接入对宿主植物植物光合作用色素含量有积极影响。随着镉浓度的升高和接触时间延长，植物光和色素含量呈现线性减少趋势。但是，在镉胁迫条件下（除了100μM，第三天样品外），接种内生菌的植物叶片中类胡萝卜素、叶绿素a和叶绿素b的含量均要高于未接菌的植株(P <0.05, MANOVAs)。另外，在同一镉浓度下，由镉胁迫导致的植物光合作用色素含量的减少量要明显小于未接菌植株，即接入内生菌的植株光合作用色素受镉影响较小。
随着营养液中重金属镉浓度的增加，植物累积了大量镉在其根部和地上部分。然而，内生菌接种与否对单位质量植物组织累积镉没有显著差异（除了茎部，镉10和100μM），即单位干重植株组织吸收的镉量相近。相比于未接菌对照植株，虽然内生菌株的植入没有极大改变重金属镉在其宿主植物组织中的浓度，但是提高了植物的生物量，由此产生的结果就是接菌的植物单株重金属镉提取量有极大提高（在10μM Cd条件下，增长量达到72±5%），这反映出内生菌的植入提高了植物提取效率。
③内生菌在降低重金属植物毒性和促进植物生长中的可能作用机制
Cd导致了植物组织中Fe含量的提高，然而Mn和Zn的含量均有所减少，除了根部Zn含量没有显著变化。这些结果表明Cd改变了其他重金属离子在植物体内的传输。接种内生菌S. nematodiphila LRE07减缓了这种由镉诱导了金属含量变化。在10、50、100μM镉浓度条件下，接种内生菌株的植株各组织（根、茎、叶）中Mn的含量都要高于未接种植株。相比于未接菌对照，接菌植物在其叶部和颈部吸收了更多的Fe和Zn。
不论接菌与否，植物组织中丙二醛含量随着镉浓度和接触时间均呈现线性提高。但是，从结果可以看出，接菌植株的丙二醛含量要低于未接菌对照，虽然两者之间没有统计性差别。另外，丙二醛含量随接触时间的增加量在接菌植株中也要小得多。
为了反映镉胁迫条件下植物体内抗氧化代谢酶的活性，我们选取了SOD,GPX, CAT和APX作为解毒活性氧毒害作用酶系的代表，测定了这些酶在镉胁迫状况下的变化。镉诱导使得SOD含量显著增加，在10μM和50μM镉浓度条件下接菌植株SOD的活性要明显要于未接种植物。CAT活性在两种处理之间没有显著区别。10μM和50μM镉条件下，接菌植物叶片中GPX和APX活性要明显高于未接菌植物。镉浓度的提高导致了两种处理的植物的根部GPX含量都呈线性增加，但是在50μM和100μM镉浓度条件下，接菌植株根部GPX含量要明显高于未接菌植株。上述结果表明接种内生菌株S. nematodiphila LRE07提高了宿主植物的抗氧化能力、降低了镉胁迫导致的活性氧伤害。
④内生菌的稳定性和在非宿主植物体内的作用
通过比较不同处理的两代植物体内内生菌菌落和植物生长状况，探讨内生菌在宿主植物体内的稳定性。结果发现，两代植物体内的目的菌株的菌落数没有显著性差异。PCR-DGGE图谱分析显示两代植株体内细菌DNA呈现较高一致性。同时传代内生菌保存对植物生长的积极影响，两代植株各项生长指数均没有显著性差异。这表明内生菌能能植物体内稳定存活，并在植物子代中保持原有活性。
将S. nematodiphila LRE07接种入豌豆并将植株置于镉胁迫条件下以探讨内生菌对非宿主植物的作用。结果发现重金属胁迫抑制植物种子萌发，内生菌接种没能缓解重金属对非宿主植物豌豆发芽率的胁迫影响。接种S. nematodiphilaLRE07能提高植物生物量，对提高植物对抗重金属胁迫有一定积极意义。但整体而言，接菌处理对非宿主植物生长无显著积极影响。
Toxic metal pollution has received considerable attention over the years as aresult of increased industrial activities. Heavy metals can be accumulated inorganisms and finally threaten human health through food chains. Therefore, aneffective and affordable solution is urgently needed to remove toxic metals fromenvironment. Established technologies to remediate metal contaminated soils arefrequently expensive and environmentally invasive. In situ phytoremediation has beenproposed as a low-cost, environmentally friendly and effective method to removetoxicant from contaminated soils. However, phytoremediation of heavy metal still hasto deal with some important shortcomings such as phytotoxicity, slower thanmechanical methods, and a limited contaminant uptake. Some phytoremediationenhancing strategies, such as transgenics and chelating agent, are comparativelyavailable but usually expensive and posing potential risks to environment. Therefore,it is important to search a microbe-strengthened phytoremediation approach toenhance plant remediation economically.
In the present study, a number of endophytes were isolated from Cdhyperaccumulator plant Solanum nigrum L.. An excellent strain Serratianematodiphila LRE07was selected to study the feasibility of phytoremediationenhancement using plant-endophyte symbiotic system. After inoculating with S.nematodiphila LRE07, plant Solanum nigrum L. was grown in hydroponics withincreasing Cd concentrations. The growth, physiological parameters, photosyntheticpigments, and Cd accumulation characteristics were investigated to study the effect ofendophyte inoculation on plant stress resistance. Further, the nutrient uptake, lipidperoxidation levels, and activated oxygen metabolism were determined order to knowthe possible mechanisms of endophyte in phytotoxicity reduction and growthpromotion involved in phytoremediation. Moreover, the stability of S. nematodiphilaLRE07and the effects of endophyte inoculation on non-host plant were investigated.The main results were indicated as follows:
①Isolation and selection of endophytic bacteria from Solanum nigrum L.
97endophytes were isolated from cadmium hyperaccumulator Solanum nigrum L.which collected at the sewage discharge canal bank of Zhuzhou Smeltery (27°52′N,113°05′E). Most of them were bacterial endophytes. All endophytes showed different degrees of heavy metals resistance by minimal inhibitory concentration test,suggesting that endophytes of hyperaccumulators are a main source which heavymetals resistant bacteria derived from. A carmine-pigmented bacterium, named asstrain LRE07, was selected from the root of Solanum nigrum L.. The isolated strainwas identified as Serratia nematodiphila by using16S rDNA analysis. S.nematodiphila LRE07showed a high degree of resistance to heavy metals, especiallyto Cu, Cd and Cr. Meanwhile, it had the ability to produce indole acetic acid andsiderophore and could solubilize mineral phosphate. The characters mentioned abovesuggested that endophytic bacterium LRE07was capable of facilitating the plantgrowth and the biomass yield. Finally, the strain LRE07was determined for the nextstep of experimental strain.
②Effect of endophyte inoculation on growth and cadmium uptake of Solanumnigrum L.
Physiological responses of S. nigrum seedings to Cd stress and the effect ofendophyte inoculation on plant stress resistance were investigated by nutrient solutionculture method. After21and70d of growth, plant samples were harvested todetermine the inoculation efficiency. Endophytic bacteria were isolated from rootsand shoots separately and plated on LB medium. The results showed that S.nematodiphila LRE07was inoculated into plant successfully. The number of CFU thatcould be re-isolated showed a time-dependent growth in roots and shoots of their hostplants and it was not inhibited by the presence of Cd. The number of re-isolated CFUtended to increase with time, indicating that S. nematodiphila LRE07could liveconcordantly with its host plants.
The increasing concentration of Cd produced growth inhibition of S. nigrum L.,measured as tillers, FW and shoot height. At0and10μM Cd, the ratio increased overtime for both infected and non-infected plants. However, at50and100μM Cd theratio rapidly reached constant values and even decreased at the end of the experiment.The higher the Cd concentration applied to the nutrient solution, the weaker the ratioof the tillers was obtained. It can be seen that the infected plants showed higher ratiovalues than noninfected controls in an equal Cd concentration as shown in Table2(P<0.05, MANOVAs), that is, greater number of leaves were obtained in infected plants.Similar beneficial effects of endophytic bacterium LRE07on plant growth wereobserved in FW, shoot height, and root weight. At10μM Cd, no visible growthinhibition was observed for infected plants and the FW of these plants increased by58%compared with non-infected ones.
With increasing Cd concentration there was a net decrease in pigment contentfrom day0to the end of the experiment whether the plants were infected or not.Endophyte-infection positively influenced the photosynthetic pigment contents of itshost plant. It can be seen that for all Cd concentrations (except for day3at100μM),chlorophyll a, chlorophyll b, and carotenoid of infected plants leaves were higher thannon-infected ones (P <0.05, MANOVAs). In addition, the decreases of photosyntheticpigment caused by Cd exposure in infected plants were much less than that of inendophyte-free plants.
Increasing concentrations of Cd in nutrient solutions led to an increase of Cdcontents in shoots and roots. Nevertheless, no significant differences were observedbetween infected and non-infected plants (except in stem: Cd10and100μM). On adry-weight basis, the plants infected and non-infected with S. nematodiphila LRE07took up approximately the same amount of cadmium. Compared with noninoculatedcontrols, endophyte inoculation did not greatly influence the cadmium concentrationsin plant tissues, but achieved a higher biomass production, thus resulting in more totalCd-uptake per plant (72±5%increase at10μM), reflecting the promotion ofphytoextraction efficiency.
③Possible mechanisms of endophyte bacterium in Cd phytotoxicity reduction andplant growth promotion.
Cd produced increase in the uptake of Fe in tissues, while the uptake of Mn andZn were decreased except no significant changes of Zn in the root. These resultsindicated that Cd induced alteration on translocation of other heavy metals in planttissues. The inoculation of endophyte S. nematodiphila LRE07alleviated thisCd-induced changes on metal contents. Mn content in infected plant tissues (roots,leaves and stems) was higher, either at10,50or100μM, than that of in non-infectedcontrols. Compared with non-symbiotic controls, symbiotic plants assimilated moreFe and Zn in leaves and stems.
The MDA content showed a linear enhancement with time and with the increasein concentration of Cd whether the plants infected or not. However, it can be seen thatthe MDA contents in infected plants were lower than that of in the noninfected plants,although differences were not statistically significant. In addition, the increment valueof MDA content with time was smaller in infected plants than those in noninfectedplants.
Activities of enzymes (SOD, GPX, CAT and APX) detoxifying the cells fromactive oxygen species were measured as representative enzymes involves in antioxidant metabolism upon Cd exposure. Cd induced obvious increase in the contentof SOD, and SOD activity is higher in infected plant tissues compared to non-infectedplants for Cd concentration of10and50μM. CAT activity in leaves showed nosignificant differences between both types of plants. Leaves of infected plants showedhigher activity of GPX and APX compared with non-symbiotic plants at10and50μM.In roots, increasing cadmium concentrations caused a linear enhancement of GPX inboth infected and non-infected plants, while for50and100μM Cd the GPX activityof infected plants were significantly higher compared with non-infected ones. Theseresults suggested that the presence of the endophytic bacterium S. nematodiphilaLRE07improved the antioxidative capability of its host plant and reduced ROS injurycaused by Cd exposure.
④Stability of endophyte in plant generation and effect of endophyte inoculationon growth and Cd resisitanse of non-host plant
Stability of endophyte in host plant generation was investigated by comparingthe endophytic bacterial conoly and plant growth condition of two different treatedplant generations. The results showed that there were no significant differencesbetween CFUs of objective strain in two plant generations. As a result of DGGEanalysis, the endophytic bacteria community composition was relatively highconsistency between the generations. Meanwhile, endophytic bacteria remained thepositive effect on plant growth, resulting in no significant differences in growthparameters between the generations. These results suggested that endophytic bacteriacould survive in plant steadily and remain its activity in plant offspring seeds.
Effect of endophyte inoculation on growth and Cd resisitanse of non-host plantwas investigated by inoculating S. nematodiphila LRE07into pea (non-host plant of S.nematodiphila LRE07) with Cd explosure. The results showed that Cd inhibited thegermination of pea seed. The endophyte inoculation could not alleviate the stresseffect in pea germination caused by Cd explosure. The inoculation of S.nematodiphila LRE07could promote plant growth under some Cd concentrations. Buton the whole, the endophyte inoculation could not significantly improve the growth ofits non-host plant.
本文自镉超累积植物龙葵中分离得到多株内生菌，通过多种性能指标筛选菌株Serratia nematodiphila LRE07作为模式菌株，用以研究利用植物-内生菌共生系统强化植物修复的可行性。将菌株S. nematodiphila LRE07接种到龙葵种子后，通过营养液培养方法深入研究了系列镉处理浓度下龙葵植株生长、生理参数、光合成色素和镉吸收累积特性，探讨了内生菌接种对植物抗压性能的影响。在此基础上，研究了镉胁迫下植物对营养元素的摄取、脂质过氧化水平和活性氧代谢的变化，以揭示内生菌在降低重金属植物毒性和促进植物生长中的可能作用机制。另外，还研究了菌株S. nematodiphila LRE07的稳定性和其在非宿主植物体内的作用效果。主要的研究内容和结果如下：
①龙葵内生菌的分离与筛选
用于内生菌分离的龙葵植株采自株洲冶炼厂（27°52′N,113°05′E）污水排水渠堤，自该植物不同组织体内分离得到97株内生菌，其中大部分为细菌。通过最小抑制浓度测定发现所有内生菌对重金属均有不同程度抗性，这表明超累积植物是筛选重金属抗性菌株的天然资源。自龙葵根部得到一株产深红色色素的细菌，命名为LRE07。通过16S rDNA分析鉴定该菌株为沙雷氏菌株（Serratianematodiphila）。S. nematodiphila LRE07表现出对多种重金属均有较高抗性，尤其是对铜、镉和铬。同时，对该菌株进行植物促生相关的生物学性能测定，结果发现S. nematodiphila LRE07具有产吲哚乙酸、铁载体和增溶矿质磷酸盐的能力。上述性能表明该菌株具有重金属抗性和植物促生潜力，因此确定S. nematodiphilaLRE07为下一步实验菌株。
②内生菌接种对龙葵植株生长和镉吸收的影响
通过营养液培养方法，研究龙葵植株对镉胁迫的响应和内生菌接种对植株抗压性能的影响。接种21天和70天后取植物样测定菌株接种效率。结果表明内生菌S. nematodiphila LRE07成功接种入植株体内，自宿主植物根部和地上部分组织内重分离得到的菌落数表现出依赖时间增长的趋势，镉胁迫未对内生菌生长造成抑制作用。上述结果表明S. nematodiphila LRE07能与宿主植物和谐共生。镉浓度的递增对龙葵生长造成了不同程度抑制作用，表现在植株的分蘖数、鲜重和株高。随着重金属接触时间延长，0μM和10μM浓度下接种植物和未接种植物的分蘖数比率均有增加，但高浓度（50和100μM）下分蘖数比率很快达到恒定值甚至在实验后期减少。营养液中重金属浓度越高，植物的分蘖数比率越低。在同一镉胁迫浓度下，接种内生菌的植株分蘖数比率要高于未接种植株（P <0.05,MANOVAs），即接种植株的叶片数更多。内生菌接种对植株生长的有利影响在植株鲜重、株高和根重上也有体现。内生菌的积极意义在低镉胁迫（10μM）下尤为显著，在该浓度下，接种植物生长未见显著抑制，其生物量比未接种植株提高了58%。
内生菌的接入对宿主植物植物光合作用色素含量有积极影响。随着镉浓度的升高和接触时间延长，植物光和色素含量呈现线性减少趋势。但是，在镉胁迫条件下（除了100μM，第三天样品外），接种内生菌的植物叶片中类胡萝卜素、叶绿素a和叶绿素b的含量均要高于未接菌的植株(P <0.05, MANOVAs)。另外，在同一镉浓度下，由镉胁迫导致的植物光合作用色素含量的减少量要明显小于未接菌植株，即接入内生菌的植株光合作用色素受镉影响较小。
随着营养液中重金属镉浓度的增加，植物累积了大量镉在其根部和地上部分。然而，内生菌接种与否对单位质量植物组织累积镉没有显著差异（除了茎部，镉10和100μM），即单位干重植株组织吸收的镉量相近。相比于未接菌对照植株，虽然内生菌株的植入没有极大改变重金属镉在其宿主植物组织中的浓度，但是提高了植物的生物量，由此产生的结果就是接菌的植物单株重金属镉提取量有极大提高（在10μM Cd条件下，增长量达到72±5%），这反映出内生菌的植入提高了植物提取效率。
③内生菌在降低重金属植物毒性和促进植物生长中的可能作用机制
Cd导致了植物组织中Fe含量的提高，然而Mn和Zn的含量均有所减少，除了根部Zn含量没有显著变化。这些结果表明Cd改变了其他重金属离子在植物体内的传输。接种内生菌S. nematodiphila LRE07减缓了这种由镉诱导了金属含量变化。在10、50、100μM镉浓度条件下，接种内生菌株的植株各组织（根、茎、叶）中Mn的含量都要高于未接种植株。相比于未接菌对照，接菌植物在其叶部和颈部吸收了更多的Fe和Zn。
不论接菌与否，植物组织中丙二醛含量随着镉浓度和接触时间均呈现线性提高。但是，从结果可以看出，接菌植株的丙二醛含量要低于未接菌对照，虽然两者之间没有统计性差别。另外，丙二醛含量随接触时间的增加量在接菌植株中也要小得多。
为了反映镉胁迫条件下植物体内抗氧化代谢酶的活性，我们选取了SOD,GPX, CAT和APX作为解毒活性氧毒害作用酶系的代表，测定了这些酶在镉胁迫状况下的变化。镉诱导使得SOD含量显著增加，在10μM和50μM镉浓度条件下接菌植株SOD的活性要明显要于未接种植物。CAT活性在两种处理之间没有显著区别。10μM和50μM镉条件下，接菌植物叶片中GPX和APX活性要明显高于未接菌植物。镉浓度的提高导致了两种处理的植物的根部GPX含量都呈线性增加，但是在50μM和100μM镉浓度条件下，接菌植株根部GPX含量要明显高于未接菌植株。上述结果表明接种内生菌株S. nematodiphila LRE07提高了宿主植物的抗氧化能力、降低了镉胁迫导致的活性氧伤害。
④内生菌的稳定性和在非宿主植物体内的作用
通过比较不同处理的两代植物体内内生菌菌落和植物生长状况，探讨内生菌在宿主植物体内的稳定性。结果发现，两代植物体内的目的菌株的菌落数没有显著性差异。PCR-DGGE图谱分析显示两代植株体内细菌DNA呈现较高一致性。同时传代内生菌保存对植物生长的积极影响，两代植株各项生长指数均没有显著性差异。这表明内生菌能能植物体内稳定存活，并在植物子代中保持原有活性。
将S. nematodiphila LRE07接种入豌豆并将植株置于镉胁迫条件下以探讨内生菌对非宿主植物的作用。结果发现重金属胁迫抑制植物种子萌发，内生菌接种没能缓解重金属对非宿主植物豌豆发芽率的胁迫影响。接种S. nematodiphilaLRE07能提高植物生物量，对提高植物对抗重金属胁迫有一定积极意义。但整体而言，接菌处理对非宿主植物生长无显著积极影响。
Toxic metal pollution has received considerable attention over the years as aresult of increased industrial activities. Heavy metals can be accumulated inorganisms and finally threaten human health through food chains. Therefore, aneffective and affordable solution is urgently needed to remove toxic metals fromenvironment. Established technologies to remediate metal contaminated soils arefrequently expensive and environmentally invasive. In situ phytoremediation has beenproposed as a low-cost, environmentally friendly and effective method to removetoxicant from contaminated soils. However, phytoremediation of heavy metal still hasto deal with some important shortcomings such as phytotoxicity, slower thanmechanical methods, and a limited contaminant uptake. Some phytoremediationenhancing strategies, such as transgenics and chelating agent, are comparativelyavailable but usually expensive and posing potential risks to environment. Therefore,it is important to search a microbe-strengthened phytoremediation approach toenhance plant remediation economically.
In the present study, a number of endophytes were isolated from Cdhyperaccumulator plant Solanum nigrum L.. An excellent strain Serratianematodiphila LRE07was selected to study the feasibility of phytoremediationenhancement using plant-endophyte symbiotic system. After inoculating with S.nematodiphila LRE07, plant Solanum nigrum L. was grown in hydroponics withincreasing Cd concentrations. The growth, physiological parameters, photosyntheticpigments, and Cd accumulation characteristics were investigated to study the effect ofendophyte inoculation on plant stress resistance. Further, the nutrient uptake, lipidperoxidation levels, and activated oxygen metabolism were determined order to knowthe possible mechanisms of endophyte in phytotoxicity reduction and growthpromotion involved in phytoremediation. Moreover, the stability of S. nematodiphilaLRE07and the effects of endophyte inoculation on non-host plant were investigated.The main results were indicated as follows:
①Isolation and selection of endophytic bacteria from Solanum nigrum L.
97endophytes were isolated from cadmium hyperaccumulator Solanum nigrum L.which collected at the sewage discharge canal bank of Zhuzhou Smeltery (27°52′N,113°05′E). Most of them were bacterial endophytes. All endophytes showed different degrees of heavy metals resistance by minimal inhibitory concentration test,suggesting that endophytes of hyperaccumulators are a main source which heavymetals resistant bacteria derived from. A carmine-pigmented bacterium, named asstrain LRE07, was selected from the root of Solanum nigrum L.. The isolated strainwas identified as Serratia nematodiphila by using16S rDNA analysis. S.nematodiphila LRE07showed a high degree of resistance to heavy metals, especiallyto Cu, Cd and Cr. Meanwhile, it had the ability to produce indole acetic acid andsiderophore and could solubilize mineral phosphate. The characters mentioned abovesuggested that endophytic bacterium LRE07was capable of facilitating the plantgrowth and the biomass yield. Finally, the strain LRE07was determined for the nextstep of experimental strain.
②Effect of endophyte inoculation on growth and cadmium uptake of Solanumnigrum L.
Physiological responses of S. nigrum seedings to Cd stress and the effect ofendophyte inoculation on plant stress resistance were investigated by nutrient solutionculture method. After21and70d of growth, plant samples were harvested todetermine the inoculation efficiency. Endophytic bacteria were isolated from rootsand shoots separately and plated on LB medium. The results showed that S.nematodiphila LRE07was inoculated into plant successfully. The number of CFU thatcould be re-isolated showed a time-dependent growth in roots and shoots of their hostplants and it was not inhibited by the presence of Cd. The number of re-isolated CFUtended to increase with time, indicating that S. nematodiphila LRE07could liveconcordantly with its host plants.
The increasing concentration of Cd produced growth inhibition of S. nigrum L.,measured as tillers, FW and shoot height. At0and10μM Cd, the ratio increased overtime for both infected and non-infected plants. However, at50and100μM Cd theratio rapidly reached constant values and even decreased at the end of the experiment.The higher the Cd concentration applied to the nutrient solution, the weaker the ratioof the tillers was obtained. It can be seen that the infected plants showed higher ratiovalues than noninfected controls in an equal Cd concentration as shown in Table2(P<0.05, MANOVAs), that is, greater number of leaves were obtained in infected plants.Similar beneficial effects of endophytic bacterium LRE07on plant growth wereobserved in FW, shoot height, and root weight. At10μM Cd, no visible growthinhibition was observed for infected plants and the FW of these plants increased by58%compared with non-infected ones.
With increasing Cd concentration there was a net decrease in pigment contentfrom day0to the end of the experiment whether the plants were infected or not.Endophyte-infection positively influenced the photosynthetic pigment contents of itshost plant. It can be seen that for all Cd concentrations (except for day3at100μM),chlorophyll a, chlorophyll b, and carotenoid of infected plants leaves were higher thannon-infected ones (P <0.05, MANOVAs). In addition, the decreases of photosyntheticpigment caused by Cd exposure in infected plants were much less than that of inendophyte-free plants.
Increasing concentrations of Cd in nutrient solutions led to an increase of Cdcontents in shoots and roots. Nevertheless, no significant differences were observedbetween infected and non-infected plants (except in stem: Cd10and100μM). On adry-weight basis, the plants infected and non-infected with S. nematodiphila LRE07took up approximately the same amount of cadmium. Compared with noninoculatedcontrols, endophyte inoculation did not greatly influence the cadmium concentrationsin plant tissues, but achieved a higher biomass production, thus resulting in more totalCd-uptake per plant (72±5%increase at10μM), reflecting the promotion ofphytoextraction efficiency.
③Possible mechanisms of endophyte bacterium in Cd phytotoxicity reduction andplant growth promotion.
Cd produced increase in the uptake of Fe in tissues, while the uptake of Mn andZn were decreased except no significant changes of Zn in the root. These resultsindicated that Cd induced alteration on translocation of other heavy metals in planttissues. The inoculation of endophyte S. nematodiphila LRE07alleviated thisCd-induced changes on metal contents. Mn content in infected plant tissues (roots,leaves and stems) was higher, either at10,50or100μM, than that of in non-infectedcontrols. Compared with non-symbiotic controls, symbiotic plants assimilated moreFe and Zn in leaves and stems.
The MDA content showed a linear enhancement with time and with the increasein concentration of Cd whether the plants infected or not. However, it can be seen thatthe MDA contents in infected plants were lower than that of in the noninfected plants,although differences were not statistically significant. In addition, the increment valueof MDA content with time was smaller in infected plants than those in noninfectedplants.
Activities of enzymes (SOD, GPX, CAT and APX) detoxifying the cells fromactive oxygen species were measured as representative enzymes involves in antioxidant metabolism upon Cd exposure. Cd induced obvious increase in the contentof SOD, and SOD activity is higher in infected plant tissues compared to non-infectedplants for Cd concentration of10and50μM. CAT activity in leaves showed nosignificant differences between both types of plants. Leaves of infected plants showedhigher activity of GPX and APX compared with non-symbiotic plants at10and50μM.In roots, increasing cadmium concentrations caused a linear enhancement of GPX inboth infected and non-infected plants, while for50and100μM Cd the GPX activityof infected plants were significantly higher compared with non-infected ones. Theseresults suggested that the presence of the endophytic bacterium S. nematodiphilaLRE07improved the antioxidative capability of its host plant and reduced ROS injurycaused by Cd exposure.
④Stability of endophyte in plant generation and effect of endophyte inoculationon growth and Cd resisitanse of non-host plant
Stability of endophyte in host plant generation was investigated by comparingthe endophytic bacterial conoly and plant growth condition of two different treatedplant generations. The results showed that there were no significant differencesbetween CFUs of objective strain in two plant generations. As a result of DGGEanalysis, the endophytic bacteria community composition was relatively highconsistency between the generations. Meanwhile, endophytic bacteria remained thepositive effect on plant growth, resulting in no significant differences in growthparameters between the generations. These results suggested that endophytic bacteriacould survive in plant steadily and remain its activity in plant offspring seeds.
Effect of endophyte inoculation on growth and Cd resisitanse of non-host plantwas investigated by inoculating S. nematodiphila LRE07into pea (non-host plant of S.nematodiphila LRE07) with Cd explosure. The results showed that Cd inhibited thegermination of pea seed. The endophyte inoculation could not alleviate the stresseffect in pea germination caused by Cd explosure. The inoculation of S.nematodiphila LRE07could promote plant growth under some Cd concentrations. Buton the whole, the endophyte inoculation could not significantly improve the growth ofits non-host plant.
引文
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