几种细菌与土壤粘粒矿物相互作用的热力学研究
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摘要
本文以高岭石、蒙脱石和针铁矿等粘粒矿物及恶臭假单胞菌(Pseudomonasputida)、枯草芽孢杆菌(Bacillus subtilis)和苏云金芽孢杆菌(Bacillus thuringlensis)为材料,研究矿物与细菌的表面吸附特征、吸附热与吸附焓变,及环境pH、离子强度和温度对吸附焓变的影响,探讨了细菌与矿物的表面吸附机理。基于实验测定的接触角和zeta电位,运用表面热力学和XDLVO理论对细菌-矿物的吸附自由能进行了理论计算。探讨了粘粒矿物对细菌代谢活性的影响,获得了细菌-粘粒矿物体系中细菌代谢的热动力学特征。研究了镉及蒙脱石-镉体系中,假单胞菌的代谢活性,从热力学角度探讨了粘土矿物影响镉毒性的机理。主要结果如下:
1.首次运用等温滴定量热技术(ITC),结合吸附、解吸等化学方法及扫描电镜技术(SEM)和傅立叶变换红外光谱技术(FTIR),揭示了细菌在高岭石、蒙脱石和针铁矿表面的主要吸附机制及其热力学特征。粘粒矿物对假单胞菌吸附率的大小顺序是:针铁矿(100%)＞高岭石(96.5%)＞蒙脱石(61.5%)。在pH等于或接近高岭石或蒙脱石的等电点时,假单胞菌在这两种矿物表面的吸附量最大,显示非静电力在吸附中的重要地位。高岭石、蒙脱石与假单胞菌的表面水分子在矿物-细菌吸附时有氢键形成,这是影响假单胞菌-矿物吸附的主要作用力之一。而表面羧基、氨基和羟基则在枯草芽胞杆菌-矿物吸附过程中起重要作用。细菌与针铁矿的吸附主要受静电力和氢键的影响。细菌与粘粒矿物的吸附均为放热反应。在相同条件下,细菌在粘粒矿物表面吸附焓变的大小顺序为:针铁矿(-7.55×10~(-8)mJ/cell)＞高岭石(-5.83×10~(-8)mJ/cell)＞蒙脱石(-4.15×10~(-8)mJ/cell),假单胞菌与粘粒矿物的吸附焓变随pH升高和离子强度增加而减小,其与层状硅酸盐矿物的吸附焓变随温度升高而增大,反应受熵与焓共同驱动,两者相互补偿。假单胞菌与针铁矿的吸附焓变随温度升高而降低,反应主要受焓驱动。枯草杆菌与矿物的吸附焓变受环境条件影响发生不规则变化,可能与革兰氏阳性的枯草芽胞杆菌细胞表面基团种类较革兰氏阴性的假单胞菌多,胞外多聚物较丰富等有关。
2.基于表面热力学和XDLVO理论,获得了粘粒矿物与细菌吸附的疏水自由能变和静电力自由能变及总自由能变。发现疏水自由能为负,显示疏水作用为引力,有利于细菌与粘粒矿物的吸附;而静电力自由能为正,表明细菌-矿物间存在静电斥力。疏水自由能显著大于静电力自由能,表明疏水作用在粘粒矿物与细菌吸附时的贡献大于静电力。两种细菌与两种矿物间的总吸附自由能为负值,意味着细菌-矿物的吸附是热力学自发过程。高岭石与细菌间的吸附自由能大于蒙脱石与细菌间的吸附自由能,表明细菌与高岭石间的亲和力较高,吸附更容易发生,这与化学吸附及滴定量热结果一致。表面热力学方法和XDLVO理论在预测细菌—矿物吸附中有重要意义,但该方法未考虑多种非DLVO效应,如细胞表面多聚物、细菌鞭毛等在吸附反应中的作用,因此还存在一定的局限性,在揭示细菌-矿物相互作用的热力学机理方面还需与其它研究技术结合。
3.首次使用等温微量热技术(IMC),研究了土壤粘粒矿物对细菌代谢活性的影响,查明了不同类型及不同浓度的土壤粘粒矿物体系中,细菌对数生长期、静止期或孢子形成期代谢活性的变化情况。发现在对数生长期,低量(5mg mL~(-1))高岭石可促进假单胞菌代谢活性,使其生长速率增大18%,代谢峰值功率提前50min出现,代谢活化能也降低2%;低量蒙脱石使枯草杆菌生长速率增加8%,代谢峰值功率提前30min出现,代谢活化能减小0.6%,显示对枯草杆菌代谢有促进效应。但不同浓度的高岭石均降低了枯草杆菌生长速率,使代谢峰值功率延迟出现且减小,代谢活化能升高,抑制枯草杆菌代谢活性。不同浓度的蒙脱石使假单胞菌生长速率减小,代谢峰值功率降低,细菌代谢活化能增大,显示对假单胞菌代谢活性有抑制作用。而针铁矿则显著抑制假单胞菌和枯草杆菌代谢活性,使它们的代谢活化能明显增加。在静止生长期,高岭石、蒙脱石和针铁矿均可抑制假单胞菌和枯草杆菌的代谢活性。对于苏云金芽孢杆菌(Bt),在高岭石、蒙脱石和针铁矿体系中,其生长速率均大于对照,总放热量也有所增加。但孢子形成期代谢峰值功率均减小,表明粘粒矿物可增强Bt对数期活性,粘粒矿物的增强效应依次为:蒙脱石＞高岭石＞针铁矿。但粘粒矿物可抑制Bt孢子形成,抑制效应是针铁矿＞高岭石＞蒙脱石。
4.运用等温微量热技术,区分了蒙脱石和无蒙脱石体系中,镉对细菌的生物毒性差异,获取了不同体系中细菌代谢的热谱参数,定量描述了蒙脱石对镉生物毒性的影响。发现在蒙脱石-镉体系中,假单胞菌代谢热功率的峰值时间、峰值功率和总放热量均显著大于无蒙脱石体系,而生长速率常数小于无蒙脱石体系。体系中蒙脱石存在与否,镉对恶臭假单胞菌的半抑制浓度(IC_(50))未发生明显变化。这些结果表明蒙脱石的存在,延长了假单胞菌生长代谢周期,明显增强了细菌的代谢活性。矿物对镉的吸附及矿物对细菌代谢活性的促进效应是矿物减轻镉毒性的重要原因。
The interaction between bacteria and mineral is one of the most important processes in soil ecosystem.Understanding the mechanisms of bacterial sorption on mineral and the influence of soil minerals on the metabolic activity is of great environmental and geological significance.In this study,kaolinite,montmorillonite and goethite were selected as model minerals and Pseudomonas putida,Bacillus subtilis and Bacillus thuringiensis were used as model bacteria.The adsorption amount,adsorption heat and adsorption enthalpy of bacteria on clay minerals were obtained by equilibrium adsorption and isothermal titration calorimetry.Based on the surface thermodynamics and XDLVO theory,the free energy changes in sorption of bacteria on minerals were calculated.The metabolic activities of bacteria in the system with the presence of various concentrations of clay minerals were monitored.The effects of minerals on bacterial activity were evaluated,and the toxicities of cadmium on P.putida in the absence and presence of montmorillonite were assessed by microcalorimetric technique.The main results were outlined as follow:
1.Isothermal titration calorimetry(ITC),with a combination of chemical adsorption-desorption,scanning electron microscopy(SEM) and Fourier transform infrared spectra(FTIR),were firstly employed to investigate the chemical characteristics and the thermodynamics of bacterial adsorption on kaolinite,montmorillonite and goethite.The greatest amount of bacteria was adsorbed by goethite(100%),followed by kaolinite(96.5%),and to a lesser extent by montmorillonite(61.5%).The maximal adsorption of P.putida on kaolinite and montmorillonite was found at pHs equivalent or close to the zero point charge of the two clay minerals,showing the important role of the non-electrostatic forces in the adsorption of bacteria.Hydrogen bonding was considered to be one of the main forces controlling the sorption of P.putida on kaolinite and montmorillonite,while the functional groups such as carboxyl,amino and hydroxyl group on surface of B.subtilis were involved in the adsorption on soil clay minerals. Electrostatic force and hydrogen bonding were the major forces governing the sorption of the bacteria on goethite.The adsorption of bacteria on soil clay minerals was exothermic process.The adsorption enthalpy of P.putida on minerals followed the order:goethite (-7.55×10~(-8) mJ/cell)＞kaolinite(-5.83×10~(-8) mJ/cell)＞montmorillonite(-4.15×10~(-8) mJ/cell).The enthalpies decreased with increasing pH and ionic strength.For kaolinite and montmorillonite,the enthalpies increased with increasing temperature,suggesting that the adsorption of P.putida was driven by entropy,and compensated by enthalpy.For goethite,the enthalpies decreased with increasing temperature,showing that the adsorption was driven by enthalpy.The enthalpies evolved in the adsorption of B.subtilis on minerals were not correlated with pH,ionic strength and temperature,probably due to the fact that more biomolecules,functional groups and extracellular polymers are present on the surface of this gram positive bacterium.
2.Based on surface thermodynamics and XDLVO theory,the free energy changes due to hydrophobic interactions(△G~H),the free energy changes originated from electrostatic force(△G~(EL)) and the total free energy changes for the adsorption of P.putida and B.subtilis on kaolinite and montmorillonite were obtained.The negative values of AGH and the positive values of△G~(EL) suggested that the hydrophobic force favored,and the electrostatic force unfavored the bacterial adsorption on kaolinite and montmorillonite. The absolute values of△G~H were much larger than those of△G~(EL),indicating that the hydrophobic interactions play a more important role than electrostatic force in bacterial adsorption.The negative total free energy changes(△G) suggested that the sorption of bacteria on clay minerals is a thermodynamically spontaneous process.Greater△G values for bacterial adsorption on kaolinite than on montmorillonite indicated higher affinity of bacteria for kaolinite than for montmorillonite,which was in agreement with the results obtained from chemical adsorption and ITC studies.
3.By using isothermal microcalorimetry,the impacts of soil clay minerals on the metabolic activity of bacteria at exponential and stationary stages were clarified.In the exponential growth stage,the growth rate constant(k) of P.putida increased by 18%,the time to reach the maximal heat power of bacterial growth was decreased by 50 min,and the metabolism activation energy was decreased by 2%in the presence of low amount(5 mg mL~(-1)) of kaolinite.Similarly,for B.subtilis,the growth rate constant increased by 8%, the peak time was decreased by 30 min,and the activation energy of bacterial metabolism was decreased by 0.2%in the presence of low amount of montmorillonite.It is assumed that the exponential growth of P.putida and B.subtilis could be stimulated by low amount of kaolinite and montmorillonite.However,the activity of B.subtilis was depressed by kaolinite,and the activity of P.putida could also be inhibited by montmorillonite.The activities of two bacteria studied were always inhibited by goethite as the presence of the mineral resulted in marked increase of metabolic activation energy of bacteria.In stationary stage,the activities of P.putida and B.subtilis were inhibited by all the minerals.For B.thuringiensis,the growth rate and total heat output were increased by the presence of the three minerals.The enhancing effect of mineral on the exponential growth was in the order of montmorillonite＞kaolinite＞goethite.The sporulation of B. thuringiensis was depressed by these minerals.The inhibitory effect followed the order: goethite＞kaolinite＞montmorillonite.
4.The toxicities of cadmium on P.putida in the absence and presence of montmorillonite were evaluated based on thermokinetic parameter.The peak time,peak height and total heat output of P.putida were increased by the presence of montmorillonite.There was no significant difference for the half inhibitory concentrations (IC_(50)) of cadmium to P.putida with and without montmorillonite.The alleviating effect of montmorillonite on the toxicity of Cd~(2+) to P.putida attributed to the adsorption of Cd~(2+) by the clay mineral and the enhancing effect of the clay on the activity of bacteria.
1.首次运用等温滴定量热技术(ITC),结合吸附、解吸等化学方法及扫描电镜技术(SEM)和傅立叶变换红外光谱技术(FTIR),揭示了细菌在高岭石、蒙脱石和针铁矿表面的主要吸附机制及其热力学特征。粘粒矿物对假单胞菌吸附率的大小顺序是:针铁矿(100%)＞高岭石(96.5%)＞蒙脱石(61.5%)。在pH等于或接近高岭石或蒙脱石的等电点时,假单胞菌在这两种矿物表面的吸附量最大,显示非静电力在吸附中的重要地位。高岭石、蒙脱石与假单胞菌的表面水分子在矿物-细菌吸附时有氢键形成,这是影响假单胞菌-矿物吸附的主要作用力之一。而表面羧基、氨基和羟基则在枯草芽胞杆菌-矿物吸附过程中起重要作用。细菌与针铁矿的吸附主要受静电力和氢键的影响。细菌与粘粒矿物的吸附均为放热反应。在相同条件下,细菌在粘粒矿物表面吸附焓变的大小顺序为:针铁矿(-7.55×10~(-8)mJ/cell)＞高岭石(-5.83×10~(-8)mJ/cell)＞蒙脱石(-4.15×10~(-8)mJ/cell),假单胞菌与粘粒矿物的吸附焓变随pH升高和离子强度增加而减小,其与层状硅酸盐矿物的吸附焓变随温度升高而增大,反应受熵与焓共同驱动,两者相互补偿。假单胞菌与针铁矿的吸附焓变随温度升高而降低,反应主要受焓驱动。枯草杆菌与矿物的吸附焓变受环境条件影响发生不规则变化,可能与革兰氏阳性的枯草芽胞杆菌细胞表面基团种类较革兰氏阴性的假单胞菌多,胞外多聚物较丰富等有关。
2.基于表面热力学和XDLVO理论,获得了粘粒矿物与细菌吸附的疏水自由能变和静电力自由能变及总自由能变。发现疏水自由能为负,显示疏水作用为引力,有利于细菌与粘粒矿物的吸附;而静电力自由能为正,表明细菌-矿物间存在静电斥力。疏水自由能显著大于静电力自由能,表明疏水作用在粘粒矿物与细菌吸附时的贡献大于静电力。两种细菌与两种矿物间的总吸附自由能为负值,意味着细菌-矿物的吸附是热力学自发过程。高岭石与细菌间的吸附自由能大于蒙脱石与细菌间的吸附自由能,表明细菌与高岭石间的亲和力较高,吸附更容易发生,这与化学吸附及滴定量热结果一致。表面热力学方法和XDLVO理论在预测细菌—矿物吸附中有重要意义,但该方法未考虑多种非DLVO效应,如细胞表面多聚物、细菌鞭毛等在吸附反应中的作用,因此还存在一定的局限性,在揭示细菌-矿物相互作用的热力学机理方面还需与其它研究技术结合。
3.首次使用等温微量热技术(IMC),研究了土壤粘粒矿物对细菌代谢活性的影响,查明了不同类型及不同浓度的土壤粘粒矿物体系中,细菌对数生长期、静止期或孢子形成期代谢活性的变化情况。发现在对数生长期,低量(5mg mL~(-1))高岭石可促进假单胞菌代谢活性,使其生长速率增大18%,代谢峰值功率提前50min出现,代谢活化能也降低2%;低量蒙脱石使枯草杆菌生长速率增加8%,代谢峰值功率提前30min出现,代谢活化能减小0.6%,显示对枯草杆菌代谢有促进效应。但不同浓度的高岭石均降低了枯草杆菌生长速率,使代谢峰值功率延迟出现且减小,代谢活化能升高,抑制枯草杆菌代谢活性。不同浓度的蒙脱石使假单胞菌生长速率减小,代谢峰值功率降低,细菌代谢活化能增大,显示对假单胞菌代谢活性有抑制作用。而针铁矿则显著抑制假单胞菌和枯草杆菌代谢活性,使它们的代谢活化能明显增加。在静止生长期,高岭石、蒙脱石和针铁矿均可抑制假单胞菌和枯草杆菌的代谢活性。对于苏云金芽孢杆菌(Bt),在高岭石、蒙脱石和针铁矿体系中,其生长速率均大于对照,总放热量也有所增加。但孢子形成期代谢峰值功率均减小,表明粘粒矿物可增强Bt对数期活性,粘粒矿物的增强效应依次为:蒙脱石＞高岭石＞针铁矿。但粘粒矿物可抑制Bt孢子形成,抑制效应是针铁矿＞高岭石＞蒙脱石。
4.运用等温微量热技术,区分了蒙脱石和无蒙脱石体系中,镉对细菌的生物毒性差异,获取了不同体系中细菌代谢的热谱参数,定量描述了蒙脱石对镉生物毒性的影响。发现在蒙脱石-镉体系中,假单胞菌代谢热功率的峰值时间、峰值功率和总放热量均显著大于无蒙脱石体系,而生长速率常数小于无蒙脱石体系。体系中蒙脱石存在与否,镉对恶臭假单胞菌的半抑制浓度(IC_(50))未发生明显变化。这些结果表明蒙脱石的存在,延长了假单胞菌生长代谢周期,明显增强了细菌的代谢活性。矿物对镉的吸附及矿物对细菌代谢活性的促进效应是矿物减轻镉毒性的重要原因。
The interaction between bacteria and mineral is one of the most important processes in soil ecosystem.Understanding the mechanisms of bacterial sorption on mineral and the influence of soil minerals on the metabolic activity is of great environmental and geological significance.In this study,kaolinite,montmorillonite and goethite were selected as model minerals and Pseudomonas putida,Bacillus subtilis and Bacillus thuringiensis were used as model bacteria.The adsorption amount,adsorption heat and adsorption enthalpy of bacteria on clay minerals were obtained by equilibrium adsorption and isothermal titration calorimetry.Based on the surface thermodynamics and XDLVO theory,the free energy changes in sorption of bacteria on minerals were calculated.The metabolic activities of bacteria in the system with the presence of various concentrations of clay minerals were monitored.The effects of minerals on bacterial activity were evaluated,and the toxicities of cadmium on P.putida in the absence and presence of montmorillonite were assessed by microcalorimetric technique.The main results were outlined as follow:
1.Isothermal titration calorimetry(ITC),with a combination of chemical adsorption-desorption,scanning electron microscopy(SEM) and Fourier transform infrared spectra(FTIR),were firstly employed to investigate the chemical characteristics and the thermodynamics of bacterial adsorption on kaolinite,montmorillonite and goethite.The greatest amount of bacteria was adsorbed by goethite(100%),followed by kaolinite(96.5%),and to a lesser extent by montmorillonite(61.5%).The maximal adsorption of P.putida on kaolinite and montmorillonite was found at pHs equivalent or close to the zero point charge of the two clay minerals,showing the important role of the non-electrostatic forces in the adsorption of bacteria.Hydrogen bonding was considered to be one of the main forces controlling the sorption of P.putida on kaolinite and montmorillonite,while the functional groups such as carboxyl,amino and hydroxyl group on surface of B.subtilis were involved in the adsorption on soil clay minerals. Electrostatic force and hydrogen bonding were the major forces governing the sorption of the bacteria on goethite.The adsorption of bacteria on soil clay minerals was exothermic process.The adsorption enthalpy of P.putida on minerals followed the order:goethite (-7.55×10~(-8) mJ/cell)＞kaolinite(-5.83×10~(-8) mJ/cell)＞montmorillonite(-4.15×10~(-8) mJ/cell).The enthalpies decreased with increasing pH and ionic strength.For kaolinite and montmorillonite,the enthalpies increased with increasing temperature,suggesting that the adsorption of P.putida was driven by entropy,and compensated by enthalpy.For goethite,the enthalpies decreased with increasing temperature,showing that the adsorption was driven by enthalpy.The enthalpies evolved in the adsorption of B.subtilis on minerals were not correlated with pH,ionic strength and temperature,probably due to the fact that more biomolecules,functional groups and extracellular polymers are present on the surface of this gram positive bacterium.
2.Based on surface thermodynamics and XDLVO theory,the free energy changes due to hydrophobic interactions(△G~H),the free energy changes originated from electrostatic force(△G~(EL)) and the total free energy changes for the adsorption of P.putida and B.subtilis on kaolinite and montmorillonite were obtained.The negative values of AGH and the positive values of△G~(EL) suggested that the hydrophobic force favored,and the electrostatic force unfavored the bacterial adsorption on kaolinite and montmorillonite. The absolute values of△G~H were much larger than those of△G~(EL),indicating that the hydrophobic interactions play a more important role than electrostatic force in bacterial adsorption.The negative total free energy changes(△G) suggested that the sorption of bacteria on clay minerals is a thermodynamically spontaneous process.Greater△G values for bacterial adsorption on kaolinite than on montmorillonite indicated higher affinity of bacteria for kaolinite than for montmorillonite,which was in agreement with the results obtained from chemical adsorption and ITC studies.
3.By using isothermal microcalorimetry,the impacts of soil clay minerals on the metabolic activity of bacteria at exponential and stationary stages were clarified.In the exponential growth stage,the growth rate constant(k) of P.putida increased by 18%,the time to reach the maximal heat power of bacterial growth was decreased by 50 min,and the metabolism activation energy was decreased by 2%in the presence of low amount(5 mg mL~(-1)) of kaolinite.Similarly,for B.subtilis,the growth rate constant increased by 8%, the peak time was decreased by 30 min,and the activation energy of bacterial metabolism was decreased by 0.2%in the presence of low amount of montmorillonite.It is assumed that the exponential growth of P.putida and B.subtilis could be stimulated by low amount of kaolinite and montmorillonite.However,the activity of B.subtilis was depressed by kaolinite,and the activity of P.putida could also be inhibited by montmorillonite.The activities of two bacteria studied were always inhibited by goethite as the presence of the mineral resulted in marked increase of metabolic activation energy of bacteria.In stationary stage,the activities of P.putida and B.subtilis were inhibited by all the minerals.For B.thuringiensis,the growth rate and total heat output were increased by the presence of the three minerals.The enhancing effect of mineral on the exponential growth was in the order of montmorillonite＞kaolinite＞goethite.The sporulation of B. thuringiensis was depressed by these minerals.The inhibitory effect followed the order: goethite＞kaolinite＞montmorillonite.
4.The toxicities of cadmium on P.putida in the absence and presence of montmorillonite were evaluated based on thermokinetic parameter.The peak time,peak height and total heat output of P.putida were increased by the presence of montmorillonite.There was no significant difference for the half inhibitory concentrations (IC_(50)) of cadmium to P.putida with and without montmorillonite.The alleviating effect of montmorillonite on the toxicity of Cd~(2+) to P.putida attributed to the adsorption of Cd~(2+) by the clay mineral and the enhancing effect of the clay on the activity of bacteria.
引文
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