丛枝菌根真菌对宝山堇菜吸收铅和镉的影响
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摘要
植物修复作为一项环境友好、成本节省的土壤重金属污染治理技术近年来广受关注，而利用土著AM真菌来提升超富集植物修复效率被认为是一种具有潜力的修复策略；因此阐明AM真菌对植物吸收重金属影响意义重大。宝山堇菜是在中国发现的为数不多的镉超富集植物之一，田间试验也证明宝山堇菜具有较强的从污染土壤中去除铅和镉的能力。本论文是首次对土著AM真菌在宝山堇菜吸收铅和镉中的作用进行了研究；研究了在不同土壤铅或镉浓度（实验一）、土壤磷水平（实验二）、时间进程（实验三）和株密度（实验四）下AM真菌对宝山堇菜铅和镉吸收的影响。本研究的主要结论如下：
1．高土壤有效铅、镉和磷浓度显著抑制宝山堇菜AM真菌侵染。施铅浓度从500mg kg~(-1)起抑制了AM真菌侵染，这种作用随铅浓度增加而增强；施镉浓度在200mg kg~(-1)时和施磷水平在500mg kg~(-1)时均显著抑制了AM菌根侵染。AM真菌侵染率随时间而提高至第9周，随后相对稳定；而株密度对AM真菌侵染率无显著影响。
2.影响AM真菌对宝山堇菜地上部磷浓度影响的主要因素是土壤磷水平，土壤有效铅镉浓度以及株密度有一定影响，而时间进程影响不显著。在矿区土中接种AM真菌均显著增加了宝山堇菜地上部磷浓度，且不受磷水平的影响。但在水稻土的施铅和施镉处理中，接种AM真菌对宝山堇菜地上部磷浓度的作用是不一致的：在实验一中无显著影响，而在实验三的Pb1000和Cd100处理中则是显著增加的。
3．土壤磷水平是影响AM真菌对宝山堇菜地上部生长作用的主要因素。在土壤磷水平较低的矿区土中，AM真菌对地上部生长的作用从不施磷处理的正面到施磷为50mg kg~(-1)和250mg kg~(-1)时的中性，再到施磷为500mg kg~(-1)时的负面。而在土壤磷水平较高的水稻土中，磷不再是宝山堇菜生长的限制因素。在施铅处理中，有效铅浓度是影响AM真菌作用的主要因素：从施铅为500mg kg~(-1)时的负面到1000和1500mg kg~(-1)的中性或正面。在施镉处理中，株密度是影响AM真菌作用的主要因素。
4.土壤中有效铅浓度是影响AM真菌对宝山堇菜地上部铅浓度作用的主要因素。在土壤DPTA提取态铅不超过221mg kg~(-1)时，AM真菌提高了宝山堇菜地上铅浓度，但在DPTA提取态铅大于或等于379mg kg~(-1)时，AM真菌降低了地上部铅浓度。未接种处理的宝山堇菜地上部铅浓度和TF值随着磷水平的提高而下降，但接种处理的所受影响不大。另外，株密度和时间进程对AM真菌的作用均无显著影响。
5.从土壤DPTA提取态镉11.5mg kg~(-1)起， AM真菌降低了宝山堇菜地上部镉浓度；且这种作用不受土壤磷水平的影响。在有效镉浓度不超过23.0mg kg~(-1)时，这种抑制作用是通过降低TF值来实现的，而在较高有效镉浓度则是通过降低根部镉浓度来实现的。株密度对AM真菌的作用有一定的影响，而时间进程则无显著影响。
本研究表明：在土壤有效铅浓度较低时AM真菌会提高宝山堇菜地上部铅浓度，但在有效铅浓度较高时则会降低地上部铅浓度。AM真菌降低了宝山堇菜地上部镉浓度，但其作用机制会因有效镉浓度高低而异。在土壤磷水平和有效铅镉浓度均较低时，AM真菌促进了宝山堇菜地上部对铅和镉的迁移总量；从而提高了对铅和镉的修复效率。在土壤有效铅或镉浓度较高时，AM真菌会降低宝山堇菜对铅和镉的吸收，这可能是AM真菌提高宝山堇菜对重金属耐性的主要机制。
Phytoextraction, as an environmentally friendly and cost-effective approach tocleaning up sites contaminated by heavy metals has received more attention in recentyears, and to employ the indigenous arbuscular mycorrhizal (AM) fungi ofhyperaccumulator species is considered as a potential strategy to enhance theefficiency of remediation. Therefore, it is of great importance to elucidate the effectsof AM fungi on plant HM uptake.
V. baoshanensis is one of the rare cadmium (Cd) hyperaccumulators found inChina and field experiment showed that V. baoshanensis had a relatively strong abilityto remove Cd and lead (Pb) from the contaminated soil. To date, it is the first time tostudy the effect of AM colonization on Cd and Pb uptake by this hyperaccumulator.The present study aims to investigate the effect of indigenous AM fungi on Cd and Pbuptake by V. baoshanensis under varying soil Cd and Pb concentrations (Exp.1), Pavailability (Exp.2), time course (Exp.3) and species density (Exp.4). Major resultsare listed as follows:
1. High availability of Pb, Cd and P significantly inhibited the AM colonization ofV. Baoshanensis. Applied Pb at500mg kg~(-1)inhibited AM colonization, and theinhibition increased with increasing soil Pb concentration, while both applied Cd at200mg kg~(-1)and applied P at500mg kg~(-1)significantly inhibited AM colonization.AM colonization increased with time till9weeks, and became relatively stablethereafter, while species density had no significant effect on AM colonization.
2. Soil P availability was the main factor affecting the role of AM fungi in shoot Pconcentration of V. Baoshanensis. Available Pb and Cd，and species density also hadcertain effect, while time course had no significant effect. In mine soils, inoculationwith AM fungi significantly increased shoot P concentration regardless of P availability. In paddy soils with applied Pb or Cd, however, the effect of AM fungi onshoot P concentration of V. baoshanensis was inconsistent: no significant effect wasobserved in Exp.1while inoculation significantly increased shoot P concentration intreatments of Pb1000and Cd100in Exp.3.
3. Soil P availability was the main factor affecting the role of AM fungi in shootgrowth of V. Baoshanensis. In mine soil with low P availability, the effect of AMfungi on shoot growth was positive without P addition, and became neutral with Paddition at50or250mg kg~(-1), and negative at applied P of500mg kg~(-1). However, inpaddy soil with high P availability, P was no longer the limiting factor of growth of V.baoshanensis. In the treatments with Pb addition, the main factor affecting the role ofAM fungi in shoot growth was available Pb: the effect was negative at Pb addition of500mg kg~(-1), and became neutral or positive at applied Pb of1000or1500mg kg~(-1). Inthe treatments with Cd addition, the main factor affecting the role of AM fungi inshoot growth was species density.
4. Soil Pb availability was the main factor affecting the role of AM fungi in shootPb concentration of V. baoshanensis. When DPTA-extractble Pb was no more than221mg kg~(-1), AM fungi increased shoot Pb concentration, while AM fungi decreasedshoot Pb concentration at DPTA-extractrable Pb no less than379mg kg~(-1). Innon-inoculated treatments, shoot Pb concentrations and TF values of V. baoshanensisdecreased with increasing P availability, while in inoculated treatments no effect wasobserved with varing P availability. In addition, species density and time course hadno significant effect on the role of AM fungi in shoot Pb concentration of V.baoshanensis.
5. AM fungi decreased shoot Cd concentration of V. baoshanensis whenDPTA-extractable Cd was no less than11.5mg kg~(-1), and the effect was regardless ofP availability. The inhibition on shoot Cd concentration was realised by decreased TFvalues at DPTA-extractable Cd no more than23.0mg kg~(-1), whereas the inhibition wasachieved via decreasing root Cd concentration at higher Cd availability. Speciesdensity had certain effect on the role of AM fungi in shoot Cd concentration while nosignificant effect was observed in time course.
The results indicated at low available Pb, AM fungi increased shoot Pbconcentration of V. baoshanensis, while at high available Pb concentration AM fungidecreased shoot Pb concentration. AM fungi decreased shoot Cd concentration of V.baoshanensis, but the mechanism of inhibition depended on Cd availability. At low available P, Pb and Cd, AM fungi increased the removal of Pb or Cd in the shoots of V.baoshanensis. At high bioavailable Pb and Cd, AM fungi decreased shoot Pb or Cdconcentration of V. baoshanensis, and this was probably the main mechanisminvolved in enhancing HMs tolerance of V. baoshanensis by AM fungi.
1．高土壤有效铅、镉和磷浓度显著抑制宝山堇菜AM真菌侵染。施铅浓度从500mg kg~(-1)起抑制了AM真菌侵染，这种作用随铅浓度增加而增强；施镉浓度在200mg kg~(-1)时和施磷水平在500mg kg~(-1)时均显著抑制了AM菌根侵染。AM真菌侵染率随时间而提高至第9周，随后相对稳定；而株密度对AM真菌侵染率无显著影响。
2.影响AM真菌对宝山堇菜地上部磷浓度影响的主要因素是土壤磷水平，土壤有效铅镉浓度以及株密度有一定影响，而时间进程影响不显著。在矿区土中接种AM真菌均显著增加了宝山堇菜地上部磷浓度，且不受磷水平的影响。但在水稻土的施铅和施镉处理中，接种AM真菌对宝山堇菜地上部磷浓度的作用是不一致的：在实验一中无显著影响，而在实验三的Pb1000和Cd100处理中则是显著增加的。
3．土壤磷水平是影响AM真菌对宝山堇菜地上部生长作用的主要因素。在土壤磷水平较低的矿区土中，AM真菌对地上部生长的作用从不施磷处理的正面到施磷为50mg kg~(-1)和250mg kg~(-1)时的中性，再到施磷为500mg kg~(-1)时的负面。而在土壤磷水平较高的水稻土中，磷不再是宝山堇菜生长的限制因素。在施铅处理中，有效铅浓度是影响AM真菌作用的主要因素：从施铅为500mg kg~(-1)时的负面到1000和1500mg kg~(-1)的中性或正面。在施镉处理中，株密度是影响AM真菌作用的主要因素。
4.土壤中有效铅浓度是影响AM真菌对宝山堇菜地上部铅浓度作用的主要因素。在土壤DPTA提取态铅不超过221mg kg~(-1)时，AM真菌提高了宝山堇菜地上铅浓度，但在DPTA提取态铅大于或等于379mg kg~(-1)时，AM真菌降低了地上部铅浓度。未接种处理的宝山堇菜地上部铅浓度和TF值随着磷水平的提高而下降，但接种处理的所受影响不大。另外，株密度和时间进程对AM真菌的作用均无显著影响。
5.从土壤DPTA提取态镉11.5mg kg~(-1)起， AM真菌降低了宝山堇菜地上部镉浓度；且这种作用不受土壤磷水平的影响。在有效镉浓度不超过23.0mg kg~(-1)时，这种抑制作用是通过降低TF值来实现的，而在较高有效镉浓度则是通过降低根部镉浓度来实现的。株密度对AM真菌的作用有一定的影响，而时间进程则无显著影响。
本研究表明：在土壤有效铅浓度较低时AM真菌会提高宝山堇菜地上部铅浓度，但在有效铅浓度较高时则会降低地上部铅浓度。AM真菌降低了宝山堇菜地上部镉浓度，但其作用机制会因有效镉浓度高低而异。在土壤磷水平和有效铅镉浓度均较低时，AM真菌促进了宝山堇菜地上部对铅和镉的迁移总量；从而提高了对铅和镉的修复效率。在土壤有效铅或镉浓度较高时，AM真菌会降低宝山堇菜对铅和镉的吸收，这可能是AM真菌提高宝山堇菜对重金属耐性的主要机制。
Phytoextraction, as an environmentally friendly and cost-effective approach tocleaning up sites contaminated by heavy metals has received more attention in recentyears, and to employ the indigenous arbuscular mycorrhizal (AM) fungi ofhyperaccumulator species is considered as a potential strategy to enhance theefficiency of remediation. Therefore, it is of great importance to elucidate the effectsof AM fungi on plant HM uptake.
V. baoshanensis is one of the rare cadmium (Cd) hyperaccumulators found inChina and field experiment showed that V. baoshanensis had a relatively strong abilityto remove Cd and lead (Pb) from the contaminated soil. To date, it is the first time tostudy the effect of AM colonization on Cd and Pb uptake by this hyperaccumulator.The present study aims to investigate the effect of indigenous AM fungi on Cd and Pbuptake by V. baoshanensis under varying soil Cd and Pb concentrations (Exp.1), Pavailability (Exp.2), time course (Exp.3) and species density (Exp.4). Major resultsare listed as follows:
1. High availability of Pb, Cd and P significantly inhibited the AM colonization ofV. Baoshanensis. Applied Pb at500mg kg~(-1)inhibited AM colonization, and theinhibition increased with increasing soil Pb concentration, while both applied Cd at200mg kg~(-1)and applied P at500mg kg~(-1)significantly inhibited AM colonization.AM colonization increased with time till9weeks, and became relatively stablethereafter, while species density had no significant effect on AM colonization.
2. Soil P availability was the main factor affecting the role of AM fungi in shoot Pconcentration of V. Baoshanensis. Available Pb and Cd，and species density also hadcertain effect, while time course had no significant effect. In mine soils, inoculationwith AM fungi significantly increased shoot P concentration regardless of P availability. In paddy soils with applied Pb or Cd, however, the effect of AM fungi onshoot P concentration of V. baoshanensis was inconsistent: no significant effect wasobserved in Exp.1while inoculation significantly increased shoot P concentration intreatments of Pb1000and Cd100in Exp.3.
3. Soil P availability was the main factor affecting the role of AM fungi in shootgrowth of V. Baoshanensis. In mine soil with low P availability, the effect of AMfungi on shoot growth was positive without P addition, and became neutral with Paddition at50or250mg kg~(-1), and negative at applied P of500mg kg~(-1). However, inpaddy soil with high P availability, P was no longer the limiting factor of growth of V.baoshanensis. In the treatments with Pb addition, the main factor affecting the role ofAM fungi in shoot growth was available Pb: the effect was negative at Pb addition of500mg kg~(-1), and became neutral or positive at applied Pb of1000or1500mg kg~(-1). Inthe treatments with Cd addition, the main factor affecting the role of AM fungi inshoot growth was species density.
4. Soil Pb availability was the main factor affecting the role of AM fungi in shootPb concentration of V. baoshanensis. When DPTA-extractble Pb was no more than221mg kg~(-1), AM fungi increased shoot Pb concentration, while AM fungi decreasedshoot Pb concentration at DPTA-extractrable Pb no less than379mg kg~(-1). Innon-inoculated treatments, shoot Pb concentrations and TF values of V. baoshanensisdecreased with increasing P availability, while in inoculated treatments no effect wasobserved with varing P availability. In addition, species density and time course hadno significant effect on the role of AM fungi in shoot Pb concentration of V.baoshanensis.
5. AM fungi decreased shoot Cd concentration of V. baoshanensis whenDPTA-extractable Cd was no less than11.5mg kg~(-1), and the effect was regardless ofP availability. The inhibition on shoot Cd concentration was realised by decreased TFvalues at DPTA-extractable Cd no more than23.0mg kg~(-1), whereas the inhibition wasachieved via decreasing root Cd concentration at higher Cd availability. Speciesdensity had certain effect on the role of AM fungi in shoot Cd concentration while nosignificant effect was observed in time course.
The results indicated at low available Pb, AM fungi increased shoot Pbconcentration of V. baoshanensis, while at high available Pb concentration AM fungidecreased shoot Pb concentration. AM fungi decreased shoot Cd concentration of V.baoshanensis, but the mechanism of inhibition depended on Cd availability. At low available P, Pb and Cd, AM fungi increased the removal of Pb or Cd in the shoots of V.baoshanensis. At high bioavailable Pb and Cd, AM fungi decreased shoot Pb or Cdconcentration of V. baoshanensis, and this was probably the main mechanisminvolved in enhancing HMs tolerance of V. baoshanensis by AM fungi.
引文
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