木霉菌REMI突变株耐受Cd机理及其对油菜富集Cd影响
摘要
多年来农田土壤重金属污染是我国农产品安全生产的主要威胁，因此研究生物修复农业土壤关键技术具有重要意义。木霉菌是广泛分布于植物根围的有益微生物，除具有防治土传植物病害的作用外，还发现可与植物形成共生体（symbiont）联合修复农化物质对环境的污染，具有修复土壤重金属污染的潜在价值。因此揭示木霉菌耐受重金属和与修复性植物互作的机理是构建木霉菌-植物联合修复农业土壤环境的技术基础。
本研究是通过限制性内切酶介导整合（Restriction enzyme mediated integration,REMI）技术构建并筛选获得耐受Cd的木霉菌突变株，研究突变株耐受Cd及其对甘蓝型油菜富集Cd的影响，主要结果如下：
以康氏木霉（Trichoderma koningii）T30为出发菌，利用REMI技术检构建了200株突变株，筛选获得10株耐受Cd突变株，与野生株（T30）相比，突变株平均耐受Cd的IC50值增加1.5~2倍。去Cd率达到23~39%，提高3~4倍。其中，突变株P6是最理想的突变株。REMI诱变技术提高了木霉菌对Cd引起的氧化胁迫效应的耐性水平。在Cd胁迫下，P6的MDA峰值小于T30，O_2~-峰值低且出现时间迟于野生株。P6的CAT被Cd胁迫诱导，活性迅速增加，4h时达峰值，增加31%。而T30仅有微小的增加。P6的SOD在4h峰值时增加45%，而T30在8h才出现峰值。P6中的GSH受Cd诱导最高增加1.6倍，T30最高增加2.3倍。
电镜观察表明：Cd主要在P6的细胞壁和液泡内富集，REMI突变可提高木霉菌细胞膜系统抗Cd损害性。P6吸附Cd后FTIR谱图发生显著变化，胞壁羟基、羧基和蛋白残基是结合Cd的主要基团。P6菌丝体在液体中吸附Cd的过程符合准二级动力学方程（R~2=0.9938），1h基本达到吸附平衡，平衡吸附量为33.2mg/g。在18~47℃的范围内温度对吸附影响不显著。吸附量随pH增加而增加。能量抑制剂叠氮化钠仅能部分抑制吸附。
通过质粒抢救和RACE技术从木霉菌中克隆了耐受Cd的相关基因，tkpah。该基因属于磷脂酶A2家族GVⅡ亚群（GV ⅡPLA2）。cDNA全长1,719bp，有一个65bp的内含子。同源重组敲除T30中的tkpah，Cd胁迫试验显示，敲除子菌丝和孢子对Cd耐性均有不同程度的提高，其中敲除子孢子对Cd和H2O2胁迫的耐受性增强最为明显。说明该基因可能负调控木霉菌孢子对Cd的耐受性。
P6菌丝蛋白质组学分析表明：Cd胁迫后的菌丝蛋白质发生明显应答反应，表现为总蛋白点数明显较少，出现16个上调两倍以上蛋白点和26个特异诱导产生的蛋白点。经MALDI-TOF-TOF MS鉴定，发现多种耐Cd胁迫蛋白如热激蛋白、磷酸甘油酸变位酶、泛素蛋白连接酶等。
盆栽试验表明：P6增强了甘蓝型油菜（沪油17和沪油20）对土壤Cd胁迫的耐受，促进油菜对土壤Cd的吸附。与CK和野生株（T30）接种处理相比，在20mgkg~(-1)Cd污染的土壤中，接种P6的沪油17地上部组织干重分别增加21%和20%，沪油20则分别增加44%和24%；在50mg kg~(-1)Cd污染的土壤中，接种P6的沪油17地上部组织干重分别增加53%和31%，沪油20则分别增加16%和17%。P6还增加了油菜地上部组织Cd浓度。与CK和T30处理相比，在20mg kg~(-1)Cd的土壤中，接种P6的沪油17地上部组织中的Cd浓度分别增加27%和13%，沪油20则分别增加8%和6%；在50mg kg~(-1)Cd的土壤中，接种P6的沪油17地上部组织中的Cd浓度分别增加35%和25%，沪油20则分别增加12%和5%。
计算净化率。与CK和T30相比，P6可以提高沪油17对20mg kg~(-1)Cd污染土壤的净化率分别为58%和35%，沪油20净化率分别提高56%和31%；对50mg kg~(-1)Cd污染土壤，接种P6的沪油17净化率分别提高110%和68%，沪油20则分别提高30%和23%。
Numerous studies demonstrate that accumulation of heavy metals in farming soil canthreaten crop production by its toxicity to plants. Also, through the food chain, heavymetals can be further accumulated in human body and eventually cause serious healthproblems. Oilseed rape (Brassica napus), widely grown in Yangtze Delta region, wasexploited in the study for remediating Cd-contaminated soil in combination ofTrichoderma koningii. To improve oilseed rape phytoextracting efficiency by Trichoderma,restriction enzymes mediation integration (REMI) was used to construct Trichodermamutants with higher Cd resistance. Of200mutants,10mutants were shown with higherCd tolerance and ability of removing Cd from growing medium.
Cd stress have been already ensured to result in more active oxygens released toinjury cell, even make its death. Based on our result, the insertional mutagenesis ofTrichoderma could lead to higher yield of ROS scaveging enzymes in mycelia than wildtype of Trichoderma, for instance, P6, a REMI mutant showed the higher activity of SOD、CAT and the higher amounts of GSH as compared with wild type of Trichoderma, whichthereby improve significantly its capability to alleviate damage to mycelia caused byover-accumulation of active oxygens in the presence of Cd stress. TEM photos showedthat Cd was mainly deposited in cell wall and vacuole of the mycelia of P6. the curves ofFTIR showed clearly that-OH,-COOH and protein in cell wall were responsible mainlyfor chelating Cd.
The kinectic curve for P6mycelia uptaking Cd conform to quadratic equation,R~2=0.9938. The maximum content of absorption was33.2mg Cd per gram mycelia. The pH of solution exerted the significant influence on the absorption of Cd by mutant P6, inshort, P6myceila could improve its power in absorption of Cd from environment as theincrease of environmental pH. The energy inhibitor NaN3was not able completely toinhibit the absorption of Cd.
A tkpah gene encoding a protein homologous to platelet activating factoracetylhydrolase isoform II [PAF-AH (II)], a member of phospholiapase A2family, wascloned by plasmid rescue from a REMI mutant strain P6of Trichoderam koningii.Thegene contained a short intron of65bp and an open reading frame of1,719bp. Thededuced amino acid sequence comprises the potential conserved GHSFG motif andcatalytic triad (Ser-272, Asp-323, His-414). Tkpah expressed in E. coli showed PAF-AHactivity. tkpah gene expression was weaker in spores than in mycelia, and wasconstitutively expressed during growth in PD media. The expressions were induced bynitrogen or carbon supplementation to minimal media and suppressed after12h treatmentwith Cd. The conidia of deletion mutants of tkpah displayed an enhanced resistance toH2O2and Cd stress, thus we inferred that the gene regulate mycelial tolerance to Cd innegative way.
By2DE analysis, we found that numbers of protein spots of P6mycelia remarkablydecreased from330to255after4h Cd treatment. Among the differential spots,26spotswere identified as unique ones, and16spots were up-regulated. MALDI-TOF-TOF MSsuccessfully identified9spots which comprised14-3-3-like protein, phosphoglycero-mutase, HSP70etc.
In pot experiment of Cd-contaminated soil, mutant P6could significantly alleviatethe negative impacts of Cd on oilseed rape (Hu17and Hu20) growth, and then improvethe Cd uptake of oilseed rape plant. At Cd-contaminated soil of20mg kg~(-1)the removal rateof Cd of Hu17combinated with P6were increased by58%and35%respectively,compared to wild type Trichoderma treatment and non-inoculated treatment(CK).Similarly the figures were56%and31%respectively for Hu20-P6combination. At Cd-contaminated soil of50mg kg~(-1), the removal rate of Cd was increased by110%and68%by Hu17-P6combination respectively, and by30%and23%respectively by Hu20-P6combination. The results suggested the Trichoderma mutant-oilseed rape symbiosissystem can be used in remediation of soil contaminated with heavy metal Cd.
本研究是通过限制性内切酶介导整合（Restriction enzyme mediated integration,REMI）技术构建并筛选获得耐受Cd的木霉菌突变株，研究突变株耐受Cd及其对甘蓝型油菜富集Cd的影响，主要结果如下：
以康氏木霉（Trichoderma koningii）T30为出发菌，利用REMI技术检构建了200株突变株，筛选获得10株耐受Cd突变株，与野生株（T30）相比，突变株平均耐受Cd的IC50值增加1.5~2倍。去Cd率达到23~39%，提高3~4倍。其中，突变株P6是最理想的突变株。REMI诱变技术提高了木霉菌对Cd引起的氧化胁迫效应的耐性水平。在Cd胁迫下，P6的MDA峰值小于T30，O_2~-峰值低且出现时间迟于野生株。P6的CAT被Cd胁迫诱导，活性迅速增加，4h时达峰值，增加31%。而T30仅有微小的增加。P6的SOD在4h峰值时增加45%，而T30在8h才出现峰值。P6中的GSH受Cd诱导最高增加1.6倍，T30最高增加2.3倍。
电镜观察表明：Cd主要在P6的细胞壁和液泡内富集，REMI突变可提高木霉菌细胞膜系统抗Cd损害性。P6吸附Cd后FTIR谱图发生显著变化，胞壁羟基、羧基和蛋白残基是结合Cd的主要基团。P6菌丝体在液体中吸附Cd的过程符合准二级动力学方程（R~2=0.9938），1h基本达到吸附平衡，平衡吸附量为33.2mg/g。在18~47℃的范围内温度对吸附影响不显著。吸附量随pH增加而增加。能量抑制剂叠氮化钠仅能部分抑制吸附。
通过质粒抢救和RACE技术从木霉菌中克隆了耐受Cd的相关基因，tkpah。该基因属于磷脂酶A2家族GVⅡ亚群（GV ⅡPLA2）。cDNA全长1,719bp，有一个65bp的内含子。同源重组敲除T30中的tkpah，Cd胁迫试验显示，敲除子菌丝和孢子对Cd耐性均有不同程度的提高，其中敲除子孢子对Cd和H2O2胁迫的耐受性增强最为明显。说明该基因可能负调控木霉菌孢子对Cd的耐受性。
P6菌丝蛋白质组学分析表明：Cd胁迫后的菌丝蛋白质发生明显应答反应，表现为总蛋白点数明显较少，出现16个上调两倍以上蛋白点和26个特异诱导产生的蛋白点。经MALDI-TOF-TOF MS鉴定，发现多种耐Cd胁迫蛋白如热激蛋白、磷酸甘油酸变位酶、泛素蛋白连接酶等。
盆栽试验表明：P6增强了甘蓝型油菜（沪油17和沪油20）对土壤Cd胁迫的耐受，促进油菜对土壤Cd的吸附。与CK和野生株（T30）接种处理相比，在20mgkg~(-1)Cd污染的土壤中，接种P6的沪油17地上部组织干重分别增加21%和20%，沪油20则分别增加44%和24%；在50mg kg~(-1)Cd污染的土壤中，接种P6的沪油17地上部组织干重分别增加53%和31%，沪油20则分别增加16%和17%。P6还增加了油菜地上部组织Cd浓度。与CK和T30处理相比，在20mg kg~(-1)Cd的土壤中，接种P6的沪油17地上部组织中的Cd浓度分别增加27%和13%，沪油20则分别增加8%和6%；在50mg kg~(-1)Cd的土壤中，接种P6的沪油17地上部组织中的Cd浓度分别增加35%和25%，沪油20则分别增加12%和5%。
计算净化率。与CK和T30相比，P6可以提高沪油17对20mg kg~(-1)Cd污染土壤的净化率分别为58%和35%，沪油20净化率分别提高56%和31%；对50mg kg~(-1)Cd污染土壤，接种P6的沪油17净化率分别提高110%和68%，沪油20则分别提高30%和23%。
Numerous studies demonstrate that accumulation of heavy metals in farming soil canthreaten crop production by its toxicity to plants. Also, through the food chain, heavymetals can be further accumulated in human body and eventually cause serious healthproblems. Oilseed rape (Brassica napus), widely grown in Yangtze Delta region, wasexploited in the study for remediating Cd-contaminated soil in combination ofTrichoderma koningii. To improve oilseed rape phytoextracting efficiency by Trichoderma,restriction enzymes mediation integration (REMI) was used to construct Trichodermamutants with higher Cd resistance. Of200mutants,10mutants were shown with higherCd tolerance and ability of removing Cd from growing medium.
Cd stress have been already ensured to result in more active oxygens released toinjury cell, even make its death. Based on our result, the insertional mutagenesis ofTrichoderma could lead to higher yield of ROS scaveging enzymes in mycelia than wildtype of Trichoderma, for instance, P6, a REMI mutant showed the higher activity of SOD、CAT and the higher amounts of GSH as compared with wild type of Trichoderma, whichthereby improve significantly its capability to alleviate damage to mycelia caused byover-accumulation of active oxygens in the presence of Cd stress. TEM photos showedthat Cd was mainly deposited in cell wall and vacuole of the mycelia of P6. the curves ofFTIR showed clearly that-OH,-COOH and protein in cell wall were responsible mainlyfor chelating Cd.
The kinectic curve for P6mycelia uptaking Cd conform to quadratic equation,R~2=0.9938. The maximum content of absorption was33.2mg Cd per gram mycelia. The pH of solution exerted the significant influence on the absorption of Cd by mutant P6, inshort, P6myceila could improve its power in absorption of Cd from environment as theincrease of environmental pH. The energy inhibitor NaN3was not able completely toinhibit the absorption of Cd.
A tkpah gene encoding a protein homologous to platelet activating factoracetylhydrolase isoform II [PAF-AH (II)], a member of phospholiapase A2family, wascloned by plasmid rescue from a REMI mutant strain P6of Trichoderam koningii.Thegene contained a short intron of65bp and an open reading frame of1,719bp. Thededuced amino acid sequence comprises the potential conserved GHSFG motif andcatalytic triad (Ser-272, Asp-323, His-414). Tkpah expressed in E. coli showed PAF-AHactivity. tkpah gene expression was weaker in spores than in mycelia, and wasconstitutively expressed during growth in PD media. The expressions were induced bynitrogen or carbon supplementation to minimal media and suppressed after12h treatmentwith Cd. The conidia of deletion mutants of tkpah displayed an enhanced resistance toH2O2and Cd stress, thus we inferred that the gene regulate mycelial tolerance to Cd innegative way.
By2DE analysis, we found that numbers of protein spots of P6mycelia remarkablydecreased from330to255after4h Cd treatment. Among the differential spots,26spotswere identified as unique ones, and16spots were up-regulated. MALDI-TOF-TOF MSsuccessfully identified9spots which comprised14-3-3-like protein, phosphoglycero-mutase, HSP70etc.
In pot experiment of Cd-contaminated soil, mutant P6could significantly alleviatethe negative impacts of Cd on oilseed rape (Hu17and Hu20) growth, and then improvethe Cd uptake of oilseed rape plant. At Cd-contaminated soil of20mg kg~(-1)the removal rateof Cd of Hu17combinated with P6were increased by58%and35%respectively,compared to wild type Trichoderma treatment and non-inoculated treatment(CK).Similarly the figures were56%and31%respectively for Hu20-P6combination. At Cd-contaminated soil of50mg kg~(-1), the removal rate of Cd was increased by110%and68%by Hu17-P6combination respectively, and by30%and23%respectively by Hu20-P6combination. The results suggested the Trichoderma mutant-oilseed rape symbiosissystem can be used in remediation of soil contaminated with heavy metal Cd.
引文
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