东南景天内生菌分离鉴定及其强化重金属超积累效应与机制
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摘要
土壤重金属污染是目前面临的一个严重的环境问题,由于采矿、冶金、金属制造业排污,农业和城市生活废物的填埋和污水灌溉,导致土壤重金属污染日益严重。植物修复是一种新型的、价格低廉并且可持续发展的原位修复技术。重金属超积累植物是修复重金属污染土壤、底泥和水体资源的良好材料。然而,大多数重金属超积累植物都存在生物量小、生长速度缓慢、根系不发达以及根际土壤重金属生物有效性低等限制其大规模修复应用的因素。为了提高植物修复效率,一些促进超积累植物生长及强化根际重金属生物有效性的策略也在逐渐引起大家的关注。
超积累生态型东南景天(Sedum alfredii Hance)是最初在浙江省衢州市古老PbZn矿址上发现的第一个中国原生的Zn/Cd超积累植物,也能积累Pb,不仅是研究植物吸收、转运和超积累Zn/Cd机理的良好模式材料,也是提升植物修复重金属污染技术的良好材料。尽管东南景天在生长速率和生物量上相比多数重金属超积累植物要有些优势,但其修复土壤重金属污染的效率还是受到生长慢、生物量小、适应性弱和土壤重金属可及性的限制,研发促进东南景天生长和抗逆、提高东南景天生物量、提高东南景天对土壤重金属可及性和吸收、转运、积累重金属能力的微生物强化技术对于发挥东南景天修复重金属污染的潜能,提高实际应用的效率至关重要。
本研究主要研究结果如下：
1.用以1-氨基环丙烷-1羧酸(ACC)为唯一氮源并添加Zn、Cd和/或Pb的多碳源无机盐基本培养基,从超积累植物东南景天原生地的植株和根际土壤中分离得到85个耐受重金属的细菌菌株。对细菌16S rRNA基因(16S rDNA)序列的分析显示所得细菌分别归属于变形菌门(55%),放线菌门(27%),厚壁菌门(11%)和拟杆菌门(7%)的20个属,其中45个(53%)菌株可能分别代表8个属的15-17个新种。从28个菌株中扩增得到ACC脱氨酶结构基因(acdS)的片段,其中21个菌株在自由培养条件下表现出ACC脱氨酶活性,另7个不表现ACC脱氨酶活性的贪Cu菌属菌株的acdS与其16S rDNA序列在系统发育上不符,可能在进化过程中通过基因水平转移获得了acdS。所有菌株都不同程度地耐受Zn、Cd和/或Pb。从根际土壤,根和地上部分离到的菌株中能耐受较高浓度Zn、Cd或Pb的比例往往与原生境中该金属的浓度正相关,显示出原生地土壤和植株中高浓度的重金属对细菌耐受重金属的选择作用和与东南景天联合细菌对生境中重金属的适应性。大多数菌株能产生吲哚乙酸,嗜铁素或溶磷,有促进植物生长和提高土壤中重金属可溶性的潜能。
2.研究了从东南景天体内分离的伯克氏菌属9Burkholderia)的SaZR4, SaMR10,鞘氨醇单胞菌属(Sphingomonas)的SaMR12和贪噬菌属(Variovorax)的SaNR1四个菌株的内生特性、在根部的定殖模式及其强化植物提取重金属的能力。激光扫描共聚焦显微镜的观测结果表明菌株SaZR4, SaMR12,和SaNR1具有在植物根部形成生物被膜(biofilm)的能力,SaZR4和SaMR12可以定殖在根内部组织。SaMR10在东南景天体内定殖的总量是最低的,对东南景天生长和提取重金属的促进作用也是最弱的。SaZR4显著强化了东南景天对Zn的提取,但是对Cd的提取没有显著影响。SaMR12和SaNR1可以在含有Zn或Cd的基质中显著促进东南景天生长并提高了对Zn或Cd的提取。结果表明所用的四株特异内生细菌能以不同的方式在植物体内定殖、调控植物生长及提取和积累重金属。SaMR12和SaNR1菌株具有强化东南景天修复Zn和Cd污染土壤的应用潜力。
3.基于伯克氏菌属的SaZR4,鞘氨醇单胞菌属的SaMR12和贪噬菌属的SaNR1能促进东南景天生长和吸收、转运Zn和/或Cd的潜能,进一步用它们接种在两种重金属复合污染土壤上生长的东南景天,检验了内生细菌对东南景天修复重金属复合污染土壤的作用；基于东南景天的多年生特性,采用两季收获地上部和多次接种,检验了内生细菌应用于强化多年生超积累植物连续提取重金属的可行性。结果表明接种内生细菌可以连续两季显著提高在两种重金属复合污染土壤上生长的东南景天地上部的生物量,提高东南景天对Zn和Cd的提取量,而且提高最后所收获根部的生物量,提高了植物对Pb和Cu的提取量,证实了连续接种内生细菌结合多季收获东南景天修复重金属复合污染土壤的可行性。
4. SaCS12是从在矿山原生境健康生长的东南景天体内分离到的一株耐受多种重金属并可以用人工培养基纯培养的真菌。根据菌落形态、孢子形态和分子鉴定初步将其确定为尖孢镰刀菌(Fusarium oxysporum)。致病性检测结果表明SaCS12对易发生枯萎病和根腐病的农作物没有致病性。通过溶液培养与土培盆栽试验研究了该真菌对东南景天的生长和吸收积累重金属的影响。水培结果表明SaCS12可以增加根长和表面积、促进侧根发育,促进了植物对营养元素的吸收和叶绿素的合成。显微观察结果表明SaCS12促进了东南景天根毛的发育,扩大了根毛区在根系中的比例。SaCS12可以促进东南景天在矿山土和水稻土两种原始重金属复合污染土壤上的生长,能够强化东南景天对土壤中重金属的活化,增加了植物对重金属的吸收及转运。和不接种对照相比,接种植物对Zn、Cd、Pb和Cu的植物修复效率在矿山土上分别增加了144、139、84和63%,在水稻土上分别增加了44、55、85和77%。研究发现了一种可培养真菌而非菌根真菌强化超积累植物修复重金属污染土壤的作用,发现了非致病尖孢镰刀菌在植物修复重金属中的应用潜力。
5.采用水培实验系统比较了内生真菌印度梨形孢(Piriformospora indica)对两种生态型东南景天生长、根系形态、营养元素吸收及重金属吸收与转运的影响。在Zn或Cd处理下,超积累生态型比非超积累生态型具有更好的耐性以及吸收与转运重金属的能力。印度梨形孢能够在两种生态型东南景天根部定殖,促进两种生态型植物根系的生长发育,提高了植物体内氮和磷含量,促进了植物生长。印度梨形孢可以提高两种生态型东南景天对Zn和Cd的耐性,但是对二者在吸收和转运重金属方面却有不同的调控作用。印度梨形孢可以提高超积累生态型对Zn和Cd的吸收及转运,提高Zn和Cd在植物地上部的积累,地上部Zn和Cd积累量分别是不接种对照的1.17-1.75倍和1.83-2.21倍。而对非超积累生态型,在200μM Zn或10μM Cd处理水平下,非超积累生态型已经表现出了重金属毒害症状。印度梨形孢可以通过降低非超积累生态型对Zn或Cd的吸收在一定程度上缓解重金属的毒害作用。研究表明印度梨形孢能强化超积累植物提取和积累重金属,在植物修复重金属污染中发挥作用,也能帮助非金属积累植物耐受重金属。
研究发现与东南景天在原生境联合生长的细菌具有丰富的多样性和新颖的基因型,有些细菌能以不同的方式在东南景天根部定殖,促进植物生长并吸收和积累重金属,并提高植物修复土壤重金属污染的效率。研究也发现了一种耐受重金属的非致病内生尖孢镰刀菌和广谱促植物生长的内生真菌印度梨形孢促进超积累植物提取和积累重金属的新功能。研究为微生物强化超积累植物东南景天修复重金属污染技术提供了实验依据和新颖的细菌和真菌资源。
Heavy metal contaminations occur in soil due to mining, anthropogenic activities and soil amendment using industrial, agricultural and municipal wastes. Phytoremediation is a novel, low-cost, sustainable technique for remediation of contaminated soil in situ. Metal hyperaccumulating plants are used for remediation of heavy metal-polluted soils, sediments, and water resources. The efficiency of phytoremediation, particularly, for large scale metal phytoextraction, is often limited by the slow growth and low aboveground biomass of the metal hyperaccumulators and the metal bioavailability in soil. However, most of hyperaccumulators grow slowly and produce shallow root systems and low shoot biomass, limiting their application for heavy metal phytoextraction. Therefore, new techniques that significantly increase metal bioavailability and accumulation and plant biomass would enhance the efficacy of phytoextraction by hyperaccumulators for future phytoremediation application.
Sedum alfredii Hance is a vegetatively propagated perennial of the Crassulaceae family and was first found in an old Pb/Zn mined area in Quzhou, Zhejiang Province, southeast of China. It is a relatively fast-growing, high-biomass Zn/Cd co-hyperaccumulator and Pb accumulator. S. alfredii has now been used as a model plant to study metal hyperaccumulation and develop new techniques to enhance phytoextraction. We aim to use indigenous plant growth-promoting microbes to enhance S. alfredii phytoextraction of heavy metals.
1. Eighty-five metal-resistant bacterial strains were isolated from rhizosphere soils and Sedum alfredii plants grown in the old Pb/Zn mining area using mineral-minimal media containing1-aminocyclopropane-l-carboxylic acid (ACC) as the sole nitrogen source and high levels of Zn, Cd and/or Pb. Analyses of bacterial16S ribosomal RNA genes (16S rDNA) revealed that the85strains were affiliated to20genera of Proteobacteria (55%), Actinobacteria (27%), Firmicutes (11%) and Bacteroidetes (7%) and that45strains (53%) might represent15-17novel species of eight genera. ACC deaminase structure genes (acdS) were amplified from28strains; ACC deaminase activities were detected from21of them in free-living states. The phylogenies of the acdS sequences from the other seven strains that did not show ACC deaminase activities were incongruent with those of their16S rDNA; these acdS genes may have evolved through horizontal transfer. All the85strains showed differential resistance to high levels of Zn, Cd and/or Pb. The percentages of the obtained bacterial strains with relatively higher metal-resistance were positively correlated to the metal concentrations of the rhizosphere soil, root and shoot tissues. This indicated the nature selection of the high level metals in the soils and plants on the bacterial metal-resistance and adaption. Most of the bacterial strains could produce indole acetic acids and siderophores, or solubilize mineral phosphate, and thus had potentials to promote plant growth and increase metal-solubility from soils.
2. Four native endophytic bacteria, Burkholderia sp. SaZR4, Burkholderia sp. SaMR10, Sphingomonas sp. SaMR12and Variovorax sp. SaNR1were used to investigate their endophytic nature and root colonization patterns and effects on phytoextraction of Zn and Cd. Laser scanning confocal microscopy revealed that gfp-tagged SaZR4, SaMR12, and SaNR1cells formed biofilms on roots and that SaZR4and SaMR12cells could invade root tissues. SaMR10showed the lowest total population associated with S. alfredii and little effect on plant growth and phytoextraction. SaZR4significantly promoted Zn-extraction but not Cd-extraction. SaMR12and SaNR1significantly promoted plant growth in substrates supplemented with Zn or Cd and phytoextraction of Zn and Cd. Together, this study have shown that the four native endophytic bacteria differently colonize the host plants and modulate metal uptake and growth of host plant, and that SaMR12and SaNR1strains are promising assistants of S. alfredii plants for phytoremediation of Zn/Cd-contaminated soil.
3. Soil experiments showed that endophytic bacteria SaZR4, SaMR12and SaNR1could increase the shoot biomass of S. alfredii in the two harvest and root biomass in the second harvest in original Pb/Zn mined soil and a multi-metal contaminated paddy soil. The phytoextraction of the four heavy metals was significantly enhanced in the two soils. Phytoextraction of Zn was approximately2-fold after inoculation with SaZR4. This study showed that continual inoculation of endophytic bacteria in the perennial Zn/Cd hyperaccumulator S. alfredii is applicable.
4. A heavy metal-resistant fungus belonged to the Fusarium oxysporum complex was isolated from the Zn/Cd co-hyperaccumulator Sedum alfredii Hance grown in a Pb/Zn mined area. This Fusarium fungus was not pathogenic to plants but promoted host growth. Hydroponic experiments showed that500μM Zn or50μM Cd combined with the fungus increased root length, branches, and surface areas, enhanced nutrient uptake and chlorophyll synthesis, leading to more vigorous hyperaccumulators with greater root systems. Soil experiments showed that the fungus increased root and shoot biomass and S. alfredii-mediated heavy metal availabilities, uptake, translocation or concentrations, and thus increased phytoextraction of Zn (144and44%), Cd (139and55%), Pb (84and85%) and Cu (63and77%) from the original Pb/Zn mined soil and a multi-metal contaminated paddy soil. Together, the nonpathogenic Fusarium fungus was able to increase S. alfredii root systems and function, metal availability and accumulation, plant biomass, and thus phytoextraction efficiency. This is the first report that showed an indigenous culturable Fusarium fungus other than mycorrhizal fungi to enhance phytoextraction by hyperaccumulators. This report showed a new avenue of microorganism-assisted phytoextraction for hyperaccumulators that are mainly non-mycorrhized Brassicaceae plants. Moreover, this report showed a new function and application potential for the worldwide distributed soil and plant-associated nonpathogenic Fusarium oxysproum fungi.
5. In hydroponic experiments, HE plants exhibited stronger tolerance and higher translocation of Zn and Cd compared with NHE. The endophytic fungus Piriformospora indica could colonize in the roots of two ecotypes and promote the root development, nitrogen and phosphorus uptake. P. indica enhanced the heavy metals tolerance of the two ecotypes. The shoots of HE accumulated1.17-1.75fold amount of Zn or1.83-2.21amount of Cd compared to the uninoculation. While for NHE, even in the treatment of200μM Zn or10μM Cd, the plant exhibited heavy metal stress and the growth was inhibited. P. indica decreased Zn or Cd uptake in the heavy metal stress conditions in the NHE. However, under the higher Zn (400μM) or Cd (20μM) treatments for NHE, both noninfested and infested plants exhibited a severe biomass reduction. The results showed that the endophytic fungus P. indica has a potential application in the phytoextraction of heavy metals by the Zn/Cd hyperaccumulator Sedum alfredii Hance. In addition, results from NHE indicated that P. indica may be used for phytostabilization of heavy metals by agricultural crops.
超积累生态型东南景天(Sedum alfredii Hance)是最初在浙江省衢州市古老PbZn矿址上发现的第一个中国原生的Zn/Cd超积累植物,也能积累Pb,不仅是研究植物吸收、转运和超积累Zn/Cd机理的良好模式材料,也是提升植物修复重金属污染技术的良好材料。尽管东南景天在生长速率和生物量上相比多数重金属超积累植物要有些优势,但其修复土壤重金属污染的效率还是受到生长慢、生物量小、适应性弱和土壤重金属可及性的限制,研发促进东南景天生长和抗逆、提高东南景天生物量、提高东南景天对土壤重金属可及性和吸收、转运、积累重金属能力的微生物强化技术对于发挥东南景天修复重金属污染的潜能,提高实际应用的效率至关重要。
本研究主要研究结果如下：
1.用以1-氨基环丙烷-1羧酸(ACC)为唯一氮源并添加Zn、Cd和/或Pb的多碳源无机盐基本培养基,从超积累植物东南景天原生地的植株和根际土壤中分离得到85个耐受重金属的细菌菌株。对细菌16S rRNA基因(16S rDNA)序列的分析显示所得细菌分别归属于变形菌门(55%),放线菌门(27%),厚壁菌门(11%)和拟杆菌门(7%)的20个属,其中45个(53%)菌株可能分别代表8个属的15-17个新种。从28个菌株中扩增得到ACC脱氨酶结构基因(acdS)的片段,其中21个菌株在自由培养条件下表现出ACC脱氨酶活性,另7个不表现ACC脱氨酶活性的贪Cu菌属菌株的acdS与其16S rDNA序列在系统发育上不符,可能在进化过程中通过基因水平转移获得了acdS。所有菌株都不同程度地耐受Zn、Cd和/或Pb。从根际土壤,根和地上部分离到的菌株中能耐受较高浓度Zn、Cd或Pb的比例往往与原生境中该金属的浓度正相关,显示出原生地土壤和植株中高浓度的重金属对细菌耐受重金属的选择作用和与东南景天联合细菌对生境中重金属的适应性。大多数菌株能产生吲哚乙酸,嗜铁素或溶磷,有促进植物生长和提高土壤中重金属可溶性的潜能。
2.研究了从东南景天体内分离的伯克氏菌属9Burkholderia)的SaZR4, SaMR10,鞘氨醇单胞菌属(Sphingomonas)的SaMR12和贪噬菌属(Variovorax)的SaNR1四个菌株的内生特性、在根部的定殖模式及其强化植物提取重金属的能力。激光扫描共聚焦显微镜的观测结果表明菌株SaZR4, SaMR12,和SaNR1具有在植物根部形成生物被膜(biofilm)的能力,SaZR4和SaMR12可以定殖在根内部组织。SaMR10在东南景天体内定殖的总量是最低的,对东南景天生长和提取重金属的促进作用也是最弱的。SaZR4显著强化了东南景天对Zn的提取,但是对Cd的提取没有显著影响。SaMR12和SaNR1可以在含有Zn或Cd的基质中显著促进东南景天生长并提高了对Zn或Cd的提取。结果表明所用的四株特异内生细菌能以不同的方式在植物体内定殖、调控植物生长及提取和积累重金属。SaMR12和SaNR1菌株具有强化东南景天修复Zn和Cd污染土壤的应用潜力。
3.基于伯克氏菌属的SaZR4,鞘氨醇单胞菌属的SaMR12和贪噬菌属的SaNR1能促进东南景天生长和吸收、转运Zn和/或Cd的潜能,进一步用它们接种在两种重金属复合污染土壤上生长的东南景天,检验了内生细菌对东南景天修复重金属复合污染土壤的作用；基于东南景天的多年生特性,采用两季收获地上部和多次接种,检验了内生细菌应用于强化多年生超积累植物连续提取重金属的可行性。结果表明接种内生细菌可以连续两季显著提高在两种重金属复合污染土壤上生长的东南景天地上部的生物量,提高东南景天对Zn和Cd的提取量,而且提高最后所收获根部的生物量,提高了植物对Pb和Cu的提取量,证实了连续接种内生细菌结合多季收获东南景天修复重金属复合污染土壤的可行性。
4. SaCS12是从在矿山原生境健康生长的东南景天体内分离到的一株耐受多种重金属并可以用人工培养基纯培养的真菌。根据菌落形态、孢子形态和分子鉴定初步将其确定为尖孢镰刀菌(Fusarium oxysporum)。致病性检测结果表明SaCS12对易发生枯萎病和根腐病的农作物没有致病性。通过溶液培养与土培盆栽试验研究了该真菌对东南景天的生长和吸收积累重金属的影响。水培结果表明SaCS12可以增加根长和表面积、促进侧根发育,促进了植物对营养元素的吸收和叶绿素的合成。显微观察结果表明SaCS12促进了东南景天根毛的发育,扩大了根毛区在根系中的比例。SaCS12可以促进东南景天在矿山土和水稻土两种原始重金属复合污染土壤上的生长,能够强化东南景天对土壤中重金属的活化,增加了植物对重金属的吸收及转运。和不接种对照相比,接种植物对Zn、Cd、Pb和Cu的植物修复效率在矿山土上分别增加了144、139、84和63%,在水稻土上分别增加了44、55、85和77%。研究发现了一种可培养真菌而非菌根真菌强化超积累植物修复重金属污染土壤的作用,发现了非致病尖孢镰刀菌在植物修复重金属中的应用潜力。
5.采用水培实验系统比较了内生真菌印度梨形孢(Piriformospora indica)对两种生态型东南景天生长、根系形态、营养元素吸收及重金属吸收与转运的影响。在Zn或Cd处理下,超积累生态型比非超积累生态型具有更好的耐性以及吸收与转运重金属的能力。印度梨形孢能够在两种生态型东南景天根部定殖,促进两种生态型植物根系的生长发育,提高了植物体内氮和磷含量,促进了植物生长。印度梨形孢可以提高两种生态型东南景天对Zn和Cd的耐性,但是对二者在吸收和转运重金属方面却有不同的调控作用。印度梨形孢可以提高超积累生态型对Zn和Cd的吸收及转运,提高Zn和Cd在植物地上部的积累,地上部Zn和Cd积累量分别是不接种对照的1.17-1.75倍和1.83-2.21倍。而对非超积累生态型,在200μM Zn或10μM Cd处理水平下,非超积累生态型已经表现出了重金属毒害症状。印度梨形孢可以通过降低非超积累生态型对Zn或Cd的吸收在一定程度上缓解重金属的毒害作用。研究表明印度梨形孢能强化超积累植物提取和积累重金属,在植物修复重金属污染中发挥作用,也能帮助非金属积累植物耐受重金属。
研究发现与东南景天在原生境联合生长的细菌具有丰富的多样性和新颖的基因型,有些细菌能以不同的方式在东南景天根部定殖,促进植物生长并吸收和积累重金属,并提高植物修复土壤重金属污染的效率。研究也发现了一种耐受重金属的非致病内生尖孢镰刀菌和广谱促植物生长的内生真菌印度梨形孢促进超积累植物提取和积累重金属的新功能。研究为微生物强化超积累植物东南景天修复重金属污染技术提供了实验依据和新颖的细菌和真菌资源。
Heavy metal contaminations occur in soil due to mining, anthropogenic activities and soil amendment using industrial, agricultural and municipal wastes. Phytoremediation is a novel, low-cost, sustainable technique for remediation of contaminated soil in situ. Metal hyperaccumulating plants are used for remediation of heavy metal-polluted soils, sediments, and water resources. The efficiency of phytoremediation, particularly, for large scale metal phytoextraction, is often limited by the slow growth and low aboveground biomass of the metal hyperaccumulators and the metal bioavailability in soil. However, most of hyperaccumulators grow slowly and produce shallow root systems and low shoot biomass, limiting their application for heavy metal phytoextraction. Therefore, new techniques that significantly increase metal bioavailability and accumulation and plant biomass would enhance the efficacy of phytoextraction by hyperaccumulators for future phytoremediation application.
Sedum alfredii Hance is a vegetatively propagated perennial of the Crassulaceae family and was first found in an old Pb/Zn mined area in Quzhou, Zhejiang Province, southeast of China. It is a relatively fast-growing, high-biomass Zn/Cd co-hyperaccumulator and Pb accumulator. S. alfredii has now been used as a model plant to study metal hyperaccumulation and develop new techniques to enhance phytoextraction. We aim to use indigenous plant growth-promoting microbes to enhance S. alfredii phytoextraction of heavy metals.
1. Eighty-five metal-resistant bacterial strains were isolated from rhizosphere soils and Sedum alfredii plants grown in the old Pb/Zn mining area using mineral-minimal media containing1-aminocyclopropane-l-carboxylic acid (ACC) as the sole nitrogen source and high levels of Zn, Cd and/or Pb. Analyses of bacterial16S ribosomal RNA genes (16S rDNA) revealed that the85strains were affiliated to20genera of Proteobacteria (55%), Actinobacteria (27%), Firmicutes (11%) and Bacteroidetes (7%) and that45strains (53%) might represent15-17novel species of eight genera. ACC deaminase structure genes (acdS) were amplified from28strains; ACC deaminase activities were detected from21of them in free-living states. The phylogenies of the acdS sequences from the other seven strains that did not show ACC deaminase activities were incongruent with those of their16S rDNA; these acdS genes may have evolved through horizontal transfer. All the85strains showed differential resistance to high levels of Zn, Cd and/or Pb. The percentages of the obtained bacterial strains with relatively higher metal-resistance were positively correlated to the metal concentrations of the rhizosphere soil, root and shoot tissues. This indicated the nature selection of the high level metals in the soils and plants on the bacterial metal-resistance and adaption. Most of the bacterial strains could produce indole acetic acids and siderophores, or solubilize mineral phosphate, and thus had potentials to promote plant growth and increase metal-solubility from soils.
2. Four native endophytic bacteria, Burkholderia sp. SaZR4, Burkholderia sp. SaMR10, Sphingomonas sp. SaMR12and Variovorax sp. SaNR1were used to investigate their endophytic nature and root colonization patterns and effects on phytoextraction of Zn and Cd. Laser scanning confocal microscopy revealed that gfp-tagged SaZR4, SaMR12, and SaNR1cells formed biofilms on roots and that SaZR4and SaMR12cells could invade root tissues. SaMR10showed the lowest total population associated with S. alfredii and little effect on plant growth and phytoextraction. SaZR4significantly promoted Zn-extraction but not Cd-extraction. SaMR12and SaNR1significantly promoted plant growth in substrates supplemented with Zn or Cd and phytoextraction of Zn and Cd. Together, this study have shown that the four native endophytic bacteria differently colonize the host plants and modulate metal uptake and growth of host plant, and that SaMR12and SaNR1strains are promising assistants of S. alfredii plants for phytoremediation of Zn/Cd-contaminated soil.
3. Soil experiments showed that endophytic bacteria SaZR4, SaMR12and SaNR1could increase the shoot biomass of S. alfredii in the two harvest and root biomass in the second harvest in original Pb/Zn mined soil and a multi-metal contaminated paddy soil. The phytoextraction of the four heavy metals was significantly enhanced in the two soils. Phytoextraction of Zn was approximately2-fold after inoculation with SaZR4. This study showed that continual inoculation of endophytic bacteria in the perennial Zn/Cd hyperaccumulator S. alfredii is applicable.
4. A heavy metal-resistant fungus belonged to the Fusarium oxysporum complex was isolated from the Zn/Cd co-hyperaccumulator Sedum alfredii Hance grown in a Pb/Zn mined area. This Fusarium fungus was not pathogenic to plants but promoted host growth. Hydroponic experiments showed that500μM Zn or50μM Cd combined with the fungus increased root length, branches, and surface areas, enhanced nutrient uptake and chlorophyll synthesis, leading to more vigorous hyperaccumulators with greater root systems. Soil experiments showed that the fungus increased root and shoot biomass and S. alfredii-mediated heavy metal availabilities, uptake, translocation or concentrations, and thus increased phytoextraction of Zn (144and44%), Cd (139and55%), Pb (84and85%) and Cu (63and77%) from the original Pb/Zn mined soil and a multi-metal contaminated paddy soil. Together, the nonpathogenic Fusarium fungus was able to increase S. alfredii root systems and function, metal availability and accumulation, plant biomass, and thus phytoextraction efficiency. This is the first report that showed an indigenous culturable Fusarium fungus other than mycorrhizal fungi to enhance phytoextraction by hyperaccumulators. This report showed a new avenue of microorganism-assisted phytoextraction for hyperaccumulators that are mainly non-mycorrhized Brassicaceae plants. Moreover, this report showed a new function and application potential for the worldwide distributed soil and plant-associated nonpathogenic Fusarium oxysproum fungi.
5. In hydroponic experiments, HE plants exhibited stronger tolerance and higher translocation of Zn and Cd compared with NHE. The endophytic fungus Piriformospora indica could colonize in the roots of two ecotypes and promote the root development, nitrogen and phosphorus uptake. P. indica enhanced the heavy metals tolerance of the two ecotypes. The shoots of HE accumulated1.17-1.75fold amount of Zn or1.83-2.21amount of Cd compared to the uninoculation. While for NHE, even in the treatment of200μM Zn or10μM Cd, the plant exhibited heavy metal stress and the growth was inhibited. P. indica decreased Zn or Cd uptake in the heavy metal stress conditions in the NHE. However, under the higher Zn (400μM) or Cd (20μM) treatments for NHE, both noninfested and infested plants exhibited a severe biomass reduction. The results showed that the endophytic fungus P. indica has a potential application in the phytoextraction of heavy metals by the Zn/Cd hyperaccumulator Sedum alfredii Hance. In addition, results from NHE indicated that P. indica may be used for phytostabilization of heavy metals by agricultural crops.
引文
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