铜耐性植物内生和根际细菌的生物多样性及其强化植物富集铜的研究
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摘要
重金属污染土壤的植物修复是一种操作简便、环境友好的新兴原位修复技术,植物体内及根际微生物可通过多种作用方式影响重金属污染环境中植物的生长和土壤重金属的形态及植物对土壤重金属的吸收,从而影响植物对重金属的修复效率。研究重金属超积累或耐性植物内生及根际细菌的生物多样性及其与植物的相互关系,不仅有助于对微生物资源及生物多样性的深入认识,丰富微生物资源库,揭示微生物多样性与植物富集重金属的关系,而且可筛选高效植物促生细菌,为重金属污染土壤植物修复特别是内生细菌强化植物联合修复重金属铜污染土壤提供理论依据和基础材料。
对不同铜污染区采集的海州香薷和鸭跖草根直接提取细菌总DNA,以799F和1492R为引物扩增部分16SrDNA片段并构建16SrDNA克隆文库,16SrDNA序列系统发育分析揭示海州香薷根部内生细菌主要包括a-, β-, y-Proteobacteria和Bacteroidetes两大类群。y-Proteobacteria在海州香薷根部细菌克隆文库中占绝对优势,分别占高、低污染区根部克隆文库的73.9%和79.0%；鸭跖草根部内生细菌主要包括a-, β-, γ-Proteobacteria、Firmicutes和Actinobacteria三大类群。属于y-Proteobacteria的克隆分别占高、低污染及无污染鸭跖草根部细菌克隆文库的41.0%、77.1%和89.9%,是本研究中检测到的最多的一大类序列,在鸭跖草根部克隆文库中占绝对优势。属于Firmicutes的克隆分别占高、低及无污染鸭跖草根部细菌克隆文库的4.8%、22.9%和10.1%。属于Actinobacteria的克隆仅在矿区高污染鸭跖草根部细菌克隆文库发现,占文库的38.6%。随着土壤铜污染程度的增加,两种植物根部y-Proteobacteria细菌数量有递减的趋势,而鸭跖草根部革兰氏阳性细菌(Firmicutes和Actinobacteria)比例逐渐增大。
通过构建16SrDNA克隆文库对铜矿废弃地海州香薷和鸭跖草根际土壤细菌群落及多样性进行研究,结果表明两种植物根际细菌群落主要可归属于10大细菌类群,其中Proteobacteria在整个克隆文库中占绝对优势。Nitrospira、Chloroflexi、 Actinobacteria是仅在海州香薷根际发现的细菌类群,Cyanobacteria是仅在鸭跖草根际发现的细菌类群。通过DGGE对不同污染区两种植物根际细菌群落的研究,发现海州香薷根际细菌主要包括四大类群：a-, β-, y-Proteobacteria、Bacteroidetes、Chloroflexi及TM7门。鸭跖草根际细菌主要包括四大类群：Acidobacteria、Bacteroidetes、a-, y-Proteobacteria、Firmicutes。植物根际细菌群落多样性可能与植物种类、重金属浓度、土壤性质等有关。随着污染程度的减轻,海州香薷根际γ-Proteobacteria细菌比例逐渐增大；鸭跖草根际细菌多样性有增加的趋势,Firmicutes所占的比例逐渐下降,而Acidobacteria所占比例逐渐增大,且在低度和无污染区逐渐出现了α-,γ-Proteobacteria。
通过平板分离法,从铜耐性植物海州香薷和鸭跖草体内和根际土壤中分离筛选出抗铜64mg.L-1的细菌62株及能以ACC为唯一氮源生长的菌株19株。利用细菌通用引物对分离筛选的供试菌株16S rDNA进行PCR扩增,获得约1500bp片段,并用限制性内切酶HaeⅢ和MspⅠ对扩增片段分别进行了限制性酶切。根据酶切聚类分析结果挑选代表性菌株测序并进行16S rDNA序列系统发育分析,结果表明铜抗性内生细菌和根际细菌具有较丰富的物种多样性。分离于海州香薷的铜抗性细菌以不动杆菌属、肠杆菌属、泛菌属等γ-Proteobacteria占优势；分离于鸭跖草的铜抗性细菌以芽孢杆菌为主的Firmicutes占优势。
通过菌株生物学特性、重金属抗性及碳酸铜溶解试验以及砂培试验初步筛选出对植物具有促生和增强铜吸收效果的3株菌株并分别鉴定为Burkholderia sp.GL12, Bacillus megaterium JL35和Sphingomonas sp.YM22。该3株菌株均能产生ACC脱氨酶、精氨酸脱羧酶、IAA和铁载体,并且能够耐受多种重金属；它们能够活化培养液中的沉淀态碱式碳酸铜。
盆栽试验表明,内生菌株GL12, JL35和YM22均能促进铜污染土壤中油菜和玉米的生长,与对照相比供试植物地上部和根部干重分别增加11-56%和48-125%。接菌处理使油菜和玉米对Cu的吸收积累量分别比对照增加31%-48%和42%-91%。
菌株GL12、JL35、YM22能在油菜和玉米根际和根内定殖,菌株JL35能在油菜和玉米叶部定殖。接菌处理的油菜和玉米根际土壤水溶性铜含量分别比对照增加16%-113%和63-94%,菌株GL12还分别使油菜和玉米根际乙酸铵提取态铜浓度增加51%和14%。接菌处理可能主要通过增强玉米根部SOD活性及ASA浓度来缓解过多重金属铜引起的过氧化伤害。玉米根际及根内生细菌群落的DGGE分析表明,与未接菌对照相比,菌株对玉米根际及根内生细菌群落有一定影响。
Phytoremediation is an emerging in-situ remediation technology due to its friendly-environment, less cost and convenient operation for remediation of heavy metal-contaminated soils. A large number of microorganisms which could colonize plant tissue interiors or rhizosphere soils, can affect growth and heavy metal accumulation of plants and metal formation in heavy metal-polluted environments by many approaches, and then improve phytoremediation efficiency. Research on endophytes and rhizobacteria diversity of hyperaccumulator or metal-tolerant plants and their interaction with plants will further understand microbial resource and biodiversity, clarify the relationship between diversity and heavy metal accumulation of plants, and can obtain and applicate plant growth promoting bacteria, offer the theoretical and experimental basis for endophytic bacteria-assisted phytoremediation of Cu-contaminated soil.
Total DNA were extracted from plant roots of Elsholtzia splendens and Commelina communis growing on different contaminated sites, partial16S rDNA were amplified with the primers799F and1492R, and then16S rDNA clone libraries were constructed. Phylogenetic analysis based on16S rDNA sequences showed that endophytic bacteria of Elsholtzia splendens roots belonged to two major groups:a-, β,γ-Proteobacteria and Bacteroidetes. Clones belonging to γ-Proteobacteria accounted for73.9%and79.0%in the high-and low-contaminated clone libraries respectively, and were predominant in clone libraries of Elsholtzia splendens roots; Endophytic bacteria of Commelina communis roots belonged to three major groups:a-,β, γ-Proteobacteria, Firmicutes and Actinobacteria. Clones belonging to y-Proteobacteria accounted for41.0%,77.1%and89.9%in the high-, low-and non-contaminated clone libraries respectively, and were predominant in clone libraries of Commelina communis roots. In addition, clones belonging to Firmicutes accounted for4.8%,22.9%and10.1%in the high-, low-and non-contaminated clone libraries of Commelina communis roots respectively.38.6%of clones affiliated with Actinobacteria were only found in the high-contaminated clone library of Commelina communis roots. An increasing abundance of gram-positive bacteria (Firmicutes and Actinobacteria) was found in Commelina communis roots, and root bacterial communities of the two plant species exhibited a decrease in y-Proteobacteria with increasing copper contamination.
Bacterial community and diversity of rhizosphere soils were studied from Elsholtzia splendens and Commelina communis growing on copper mine wasteland by16S rDNA clone libraries, the results showed that rhizobacterial communities from the two plant species were affiliated with ten major groups. Proteobacteria were the most abundant group in the clone libraries. Nitrospira, Chloroflexi and Actinobacteria were only found in the rhizosphere soil of Elsholtzia splendens, and Cyanobacteria were only found in Commelina communis. Study on bacterial community of rhizosphere soils from the two plant species in different contaminated sites by DGGE found that rhizosphere bacteria from Elsholtzia splendens belonged to four major groups:a-, β-, γ-Proteobacteria, Bacteroidetes, Chloroflexi and TM7; Rhizosphere bacteria from Commelina communis belonged to four major groups:Acidobacteria, Bacteroidetes, a-, y-Proteobacteria and Firmicutes. Community composition of rhizosphere bacteria may be related to plant species, metal concentration and soil conditions. As copper contamination decreased, an increasing abundance of y-Proteobacteria was found in Elsholtzia splendens rhizosphere, rhizosphere communities of Commelina communis exhibited a decrease in Firmicutes, but an increase in Acidobacteria, and a-, γ-Proteobacteria appeared in low-, non-contaminated sites.
Sixty two strains that were resistant to Cu (64mg.L-1) and19strains that can grow on ACC as the sole N source were screened out from plant tissue interiors and rhizosphere soils of copper-tolerant plant species Elsholtzia splendens and Commelina communis by conventional plate culture technique.16S rDNA of the tested strains were amplified by PCR with the universal bacterial primers, and were digested with the restriction endonucleases Msp I and Hae III respectively.16S rDNA sequences phylogenetic analysis of representative strains with diverse ARDRA patterns indicated that the Cu-resistant-endophytic bacteria and rhizobacteria showed abundant diversity. γ-Proteobacteria (mainly Acinetobacter, Enterobacter and Pantoea) was predominant in Cu-resistant bactaria isolates from Elsholtzia splendens; Firmicutes (mainly Bacillus) was predominant in Cu-resistant bactaria isolates from Commelina communis.
Three strains with plant growth promotioning characteristics, heavy metal resistance and copper solubilization were screened out based on plant growth promotion and improvement of copper uptake of the plant and identified as Burkholderia sp.GL12, Bacillus megaterium JL35, Sphingomonas sp.YM22. Strains GL12, JL35and YM22could produce ACC deaminase, arginine decarboxylase, IAA, siderophore, and be resistant to metals; In the Cu2(OH)2CO3solubilization experiment, inoculation with the strains was found to increase the Cu2+concentration compared to the controls.
The effects of Strains GL12, JL35and YM22on plant growth and Cu uptake were studied by pot experiment. The results showed that the inoculation with tested endophytic bacteria GL12, JL35and YM22could promote rape and maize growth in Cu-contaminated soils, and increase dry weights by11-56%in shoots and by48-125%in roots compared to the controls. The Cu uptakes of inoculated rape and maize plants were increased from31%to48%and from42%to91%compared to the uninoculated control, respectively.
Strains GL12, JL35and YM22could colonize in rhizosphere and root interiors of the rape and maize, and strain JL35could colonize in leaves of rape and maize. The water-extracted Cu concentration of rhizosphere soils of the inoculated rape and maize plants was increased from16%to113%and from63%to94%compared to the controls, respectively. Moreover, the inoculation with strain GL12was found to significantly increase NH4OAc-extracted Cu concentration by51%for rape and by14%for maize, compared to the controls, respectively. The inoculation with the strains can alleviate peroxidation level under copper stress by the improvement of SOD activity and ASA concentration in maize roots. Furthermore, the research on rhizosphere bacterial and root endophytic bacterial community from maize by DGGE showed that there were some influences of the inoculation with strains on rhizosphere bacterial and root endophytic bacterial community compared to the uninoculated control.
对不同铜污染区采集的海州香薷和鸭跖草根直接提取细菌总DNA,以799F和1492R为引物扩增部分16SrDNA片段并构建16SrDNA克隆文库,16SrDNA序列系统发育分析揭示海州香薷根部内生细菌主要包括a-, β-, y-Proteobacteria和Bacteroidetes两大类群。y-Proteobacteria在海州香薷根部细菌克隆文库中占绝对优势,分别占高、低污染区根部克隆文库的73.9%和79.0%；鸭跖草根部内生细菌主要包括a-, β-, γ-Proteobacteria、Firmicutes和Actinobacteria三大类群。属于y-Proteobacteria的克隆分别占高、低污染及无污染鸭跖草根部细菌克隆文库的41.0%、77.1%和89.9%,是本研究中检测到的最多的一大类序列,在鸭跖草根部克隆文库中占绝对优势。属于Firmicutes的克隆分别占高、低及无污染鸭跖草根部细菌克隆文库的4.8%、22.9%和10.1%。属于Actinobacteria的克隆仅在矿区高污染鸭跖草根部细菌克隆文库发现,占文库的38.6%。随着土壤铜污染程度的增加,两种植物根部y-Proteobacteria细菌数量有递减的趋势,而鸭跖草根部革兰氏阳性细菌(Firmicutes和Actinobacteria)比例逐渐增大。
通过构建16SrDNA克隆文库对铜矿废弃地海州香薷和鸭跖草根际土壤细菌群落及多样性进行研究,结果表明两种植物根际细菌群落主要可归属于10大细菌类群,其中Proteobacteria在整个克隆文库中占绝对优势。Nitrospira、Chloroflexi、 Actinobacteria是仅在海州香薷根际发现的细菌类群,Cyanobacteria是仅在鸭跖草根际发现的细菌类群。通过DGGE对不同污染区两种植物根际细菌群落的研究,发现海州香薷根际细菌主要包括四大类群：a-, β-, y-Proteobacteria、Bacteroidetes、Chloroflexi及TM7门。鸭跖草根际细菌主要包括四大类群：Acidobacteria、Bacteroidetes、a-, y-Proteobacteria、Firmicutes。植物根际细菌群落多样性可能与植物种类、重金属浓度、土壤性质等有关。随着污染程度的减轻,海州香薷根际γ-Proteobacteria细菌比例逐渐增大；鸭跖草根际细菌多样性有增加的趋势,Firmicutes所占的比例逐渐下降,而Acidobacteria所占比例逐渐增大,且在低度和无污染区逐渐出现了α-,γ-Proteobacteria。
通过平板分离法,从铜耐性植物海州香薷和鸭跖草体内和根际土壤中分离筛选出抗铜64mg.L-1的细菌62株及能以ACC为唯一氮源生长的菌株19株。利用细菌通用引物对分离筛选的供试菌株16S rDNA进行PCR扩增,获得约1500bp片段,并用限制性内切酶HaeⅢ和MspⅠ对扩增片段分别进行了限制性酶切。根据酶切聚类分析结果挑选代表性菌株测序并进行16S rDNA序列系统发育分析,结果表明铜抗性内生细菌和根际细菌具有较丰富的物种多样性。分离于海州香薷的铜抗性细菌以不动杆菌属、肠杆菌属、泛菌属等γ-Proteobacteria占优势；分离于鸭跖草的铜抗性细菌以芽孢杆菌为主的Firmicutes占优势。
通过菌株生物学特性、重金属抗性及碳酸铜溶解试验以及砂培试验初步筛选出对植物具有促生和增强铜吸收效果的3株菌株并分别鉴定为Burkholderia sp.GL12, Bacillus megaterium JL35和Sphingomonas sp.YM22。该3株菌株均能产生ACC脱氨酶、精氨酸脱羧酶、IAA和铁载体,并且能够耐受多种重金属；它们能够活化培养液中的沉淀态碱式碳酸铜。
盆栽试验表明,内生菌株GL12, JL35和YM22均能促进铜污染土壤中油菜和玉米的生长,与对照相比供试植物地上部和根部干重分别增加11-56%和48-125%。接菌处理使油菜和玉米对Cu的吸收积累量分别比对照增加31%-48%和42%-91%。
菌株GL12、JL35、YM22能在油菜和玉米根际和根内定殖,菌株JL35能在油菜和玉米叶部定殖。接菌处理的油菜和玉米根际土壤水溶性铜含量分别比对照增加16%-113%和63-94%,菌株GL12还分别使油菜和玉米根际乙酸铵提取态铜浓度增加51%和14%。接菌处理可能主要通过增强玉米根部SOD活性及ASA浓度来缓解过多重金属铜引起的过氧化伤害。玉米根际及根内生细菌群落的DGGE分析表明,与未接菌对照相比,菌株对玉米根际及根内生细菌群落有一定影响。
Phytoremediation is an emerging in-situ remediation technology due to its friendly-environment, less cost and convenient operation for remediation of heavy metal-contaminated soils. A large number of microorganisms which could colonize plant tissue interiors or rhizosphere soils, can affect growth and heavy metal accumulation of plants and metal formation in heavy metal-polluted environments by many approaches, and then improve phytoremediation efficiency. Research on endophytes and rhizobacteria diversity of hyperaccumulator or metal-tolerant plants and their interaction with plants will further understand microbial resource and biodiversity, clarify the relationship between diversity and heavy metal accumulation of plants, and can obtain and applicate plant growth promoting bacteria, offer the theoretical and experimental basis for endophytic bacteria-assisted phytoremediation of Cu-contaminated soil.
Total DNA were extracted from plant roots of Elsholtzia splendens and Commelina communis growing on different contaminated sites, partial16S rDNA were amplified with the primers799F and1492R, and then16S rDNA clone libraries were constructed. Phylogenetic analysis based on16S rDNA sequences showed that endophytic bacteria of Elsholtzia splendens roots belonged to two major groups:a-, β,γ-Proteobacteria and Bacteroidetes. Clones belonging to γ-Proteobacteria accounted for73.9%and79.0%in the high-and low-contaminated clone libraries respectively, and were predominant in clone libraries of Elsholtzia splendens roots; Endophytic bacteria of Commelina communis roots belonged to three major groups:a-,β, γ-Proteobacteria, Firmicutes and Actinobacteria. Clones belonging to y-Proteobacteria accounted for41.0%,77.1%and89.9%in the high-, low-and non-contaminated clone libraries respectively, and were predominant in clone libraries of Commelina communis roots. In addition, clones belonging to Firmicutes accounted for4.8%,22.9%and10.1%in the high-, low-and non-contaminated clone libraries of Commelina communis roots respectively.38.6%of clones affiliated with Actinobacteria were only found in the high-contaminated clone library of Commelina communis roots. An increasing abundance of gram-positive bacteria (Firmicutes and Actinobacteria) was found in Commelina communis roots, and root bacterial communities of the two plant species exhibited a decrease in y-Proteobacteria with increasing copper contamination.
Bacterial community and diversity of rhizosphere soils were studied from Elsholtzia splendens and Commelina communis growing on copper mine wasteland by16S rDNA clone libraries, the results showed that rhizobacterial communities from the two plant species were affiliated with ten major groups. Proteobacteria were the most abundant group in the clone libraries. Nitrospira, Chloroflexi and Actinobacteria were only found in the rhizosphere soil of Elsholtzia splendens, and Cyanobacteria were only found in Commelina communis. Study on bacterial community of rhizosphere soils from the two plant species in different contaminated sites by DGGE found that rhizosphere bacteria from Elsholtzia splendens belonged to four major groups:a-, β-, γ-Proteobacteria, Bacteroidetes, Chloroflexi and TM7; Rhizosphere bacteria from Commelina communis belonged to four major groups:Acidobacteria, Bacteroidetes, a-, y-Proteobacteria and Firmicutes. Community composition of rhizosphere bacteria may be related to plant species, metal concentration and soil conditions. As copper contamination decreased, an increasing abundance of y-Proteobacteria was found in Elsholtzia splendens rhizosphere, rhizosphere communities of Commelina communis exhibited a decrease in Firmicutes, but an increase in Acidobacteria, and a-, γ-Proteobacteria appeared in low-, non-contaminated sites.
Sixty two strains that were resistant to Cu (64mg.L-1) and19strains that can grow on ACC as the sole N source were screened out from plant tissue interiors and rhizosphere soils of copper-tolerant plant species Elsholtzia splendens and Commelina communis by conventional plate culture technique.16S rDNA of the tested strains were amplified by PCR with the universal bacterial primers, and were digested with the restriction endonucleases Msp I and Hae III respectively.16S rDNA sequences phylogenetic analysis of representative strains with diverse ARDRA patterns indicated that the Cu-resistant-endophytic bacteria and rhizobacteria showed abundant diversity. γ-Proteobacteria (mainly Acinetobacter, Enterobacter and Pantoea) was predominant in Cu-resistant bactaria isolates from Elsholtzia splendens; Firmicutes (mainly Bacillus) was predominant in Cu-resistant bactaria isolates from Commelina communis.
Three strains with plant growth promotioning characteristics, heavy metal resistance and copper solubilization were screened out based on plant growth promotion and improvement of copper uptake of the plant and identified as Burkholderia sp.GL12, Bacillus megaterium JL35, Sphingomonas sp.YM22. Strains GL12, JL35and YM22could produce ACC deaminase, arginine decarboxylase, IAA, siderophore, and be resistant to metals; In the Cu2(OH)2CO3solubilization experiment, inoculation with the strains was found to increase the Cu2+concentration compared to the controls.
The effects of Strains GL12, JL35and YM22on plant growth and Cu uptake were studied by pot experiment. The results showed that the inoculation with tested endophytic bacteria GL12, JL35and YM22could promote rape and maize growth in Cu-contaminated soils, and increase dry weights by11-56%in shoots and by48-125%in roots compared to the controls. The Cu uptakes of inoculated rape and maize plants were increased from31%to48%and from42%to91%compared to the uninoculated control, respectively.
Strains GL12, JL35and YM22could colonize in rhizosphere and root interiors of the rape and maize, and strain JL35could colonize in leaves of rape and maize. The water-extracted Cu concentration of rhizosphere soils of the inoculated rape and maize plants was increased from16%to113%and from63%to94%compared to the controls, respectively. Moreover, the inoculation with strain GL12was found to significantly increase NH4OAc-extracted Cu concentration by51%for rape and by14%for maize, compared to the controls, respectively. The inoculation with the strains can alleviate peroxidation level under copper stress by the improvement of SOD activity and ASA concentration in maize roots. Furthermore, the research on rhizosphere bacterial and root endophytic bacterial community from maize by DGGE showed that there were some influences of the inoculation with strains on rhizosphere bacterial and root endophytic bacterial community compared to the uninoculated control.
引文
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