Bt毒素在矿物和土壤胶体上的吸附和残留研究
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摘要
转Bt基因作物发展迅速,其释放的毒素可能在土壤中积累,影响土壤生态系统的功能和生物多样性,引发环境安全问题。本文研究了Bt毒素在矿物(蒙脱石、高岭石、针铁矿和二氧化硅)和土壤(红壤、砖红壤、黄棕壤和黄褐土)胶体表面的吸附特性,用动力学和热力学方法探讨了Bt毒素的吸附规律和时间、温度、pH、低分子量有机酸盐、无机配体等因素对吸附的影响,并用红外光谱、圆二色谱和荧光光谱等观察了供试胶体吸附对Bt毒素结构的影响,剖析了被吸附Bt毒素的解吸规律,还研究了在有或无外源微生物条件下土壤中外加Bt毒素的残留特性。本研究将土壤化学与现代生物技术相融合,为转Bt基因植物代谢产物的环境效应及生态安全评价提供参考。主要结果如下:
1)Bt毒素在供试胶体表面的等温吸附曲线为L型,可用Langmuir和Freundlich方程拟合,且Freundlich方程拟合效果更好。在Bt毒素初始浓度为0-1 000 mg L~(-1)时,其在矿物胶体表面的吸附量随温度升高而减少,在供试胶体表面吸附的标准自由能(Δ_rG_m~θ)为负值,表明该吸附是自发反应。Bt毒素除在蒙脱石表面吸附的标准焓变(Δ_rH_m~θ)为正值是吸热反应外,在其它供试胶体表面吸附的标准焓变均为负值,是放热反应;且所有吸附的标准焓变值均小于40 kJ mol~(-1),说明该吸附主要是物理吸附。
2)Bt毒素在供试胶体表面能快速吸附,0.25 h的吸附量约占平衡吸附量的64%-80%,1-2 h基本达平衡。该吸附可用粒子内扩散模型拟合,常数C值不为0,表明粒子内扩散速度不是Bt毒素在供试胶体表面吸附快慢的唯一控制因素。此外,其吸附动力学也符合一级方程、二级方程和Elovich方程,以二级动力学方程的拟合程度最好,说明该吸附主要受表面吸附控制。
3)针铁矿、蒙脱石、高岭石、黄棕壤和黄褐土等胶体在pH 6时对Bt毒素的吸附量最大,二氧化硅、红壤和砖红壤等胶体在pH 7时吸附量最大,其吸附量一般随pH的升高而降低,这主要与Bt毒素的等电点及供试胶体的电荷零点有关。
4)低分子量有机酸盐和无机配体对Bt毒素在供试胶体表面的吸附受的影响。对Bt毒素的吸附低浓度(<10 mmol L~(-1))有机酸盐(乙酸盐、草酸盐、柠檬酸盐)起抑制作用,高浓度有机酸盐起促进作用;不同有机酸盐抑制作用的强弱为草酸盐<柠檬酸盐≤乙酸盐。无机配体对Bt毒素吸附的影响与有机酸盐相反,低浓度无机盐促进其吸附,高浓度的起抑制作用,不同离子的影响为H_2PO_4~->NO_3~-,NH_4~+>K~+。
5)吸附态Bt毒素很难被解吸。用0.1 mmol L~(-1) NaCl作解吸剂时,供试胶体表面吸附态Bt毒素的解吸率<5.3%,磷酸缓冲液作解吸剂时其解吸率<13.1%,表明它只有少部分以静电和配位的方式吸附在供试胶体表面。在有低分子量有机酸盐等配体存在条件下,NaCl和磷酸缓冲液对吸附态Bt毒素的解吸率均有显著增加(除乙酸盐体系中蒙脱石和砖红壤吸附态Bt毒素外),表明有机酸盐的存在一定程度上增加了静电和配位吸附,影响大小为草酸盐>柠檬酸盐>乙酸盐。
6)荧光光谱显示Bt毒素在282 nm激发光的作用下有酪氨酸(314.5 nm)和色氨酸(338 nm)荧光峰,其与高岭石、蒙脱石及土壤等胶体作用后色氨酸荧光峰红移了5-9.5 nm,酪氨酸荧光峰没有变化;与针铁矿和二氧化硅胶体作用后色氨酸和酪氨酸的荧光峰蓝移了0.5-1 nm,变化不明显。荧光峰的红移表明Bt毒素的吸附导致其内源荧光基团所处的分子环境极性增强。圆二色光谱表明吸附导致Bt毒素的二级结构改变,α螺旋和β折叠含量增加,而β转角和无规则卷曲含量降低。红外光谱显示,吸附后Bt毒素的酰胺Ⅰ带(1653 cm~(-1))和酰胺Ⅱ带(1543 cm~(-1))等特征光谱峰未发生明显位移,只是1055,1240和2362 cm~(-1)处的峰消失,表明C-N残基在供试胶体吸附Bt毒素中起重要作用,且吸附后毒素的二级结构变化不明显。
7)25℃条件下,初始浓度为0.3μg g~(-1)的游离态Bt毒素随时间增加残留量逐渐减少,10天时残留量小于原始加入量的55%,培养40天后约降解了80%,表明游离态Bt毒素降解速率较快。有外源微生物存在,其降解速率更快,表明微生物能利用游离态Bt毒素;双指数模型拟合结果表明,微生物存在下其降解速率更快,且从种植非转Bt植物上壤中提取的微生物对游离态Bt毒素的利用更快。Bt毒素在供试灭菌土壤(砖红壤、黄褐土和黄棕壤)中的残留量随着培养时间的推移先增加(1-2天)后逐步减少,降解50%的时间(DT_(50))为10天左右,外源微生物对土壤中Bt毒素的降解速率影响不明显。初始阶段Bt毒素在3种土壤中降解速度的差异不明显,而后期在黄棕壤和黄褐土中的降解速率比在砖红壤中的慢。
The wide-spread cultivation of transgenic plants would pose a risk to natural and agricultural ecosystems, especially the transgenic Bacillus thuringinesis (Bt) plants, for they release Bt toxin to the soil ecosystem by root exudates, plant biomass and pollen. This study investigated the adsorption and desorption of Bt toxin by montmorillinite, kaolinite, goethite, silicon dioxide, red soil (Ultisol), latosol (Oxisol), yellow brown soil (Alfisol) and yellow cinnamon soil (Alfisol). The kinetic and thermodynamic parameters were analyzed to understand the adsorption mechanisms, while infrared spectra, circular dichroism (CD) spectra and fluorescence spectra were used to investigate the secondary structure changes of Bt toxin during the adsorption and desorption process. The remaining of Bt toxin in soils with or without microorganisms were also studied. This investigation helps to evaluate the behavior and fate of Bt toxins in the soil ecosystem. The main results are described below.
1) The isotherm adsorption curves were L-type, and the adsorption data fitted well with both Langmuir and Freundlich isotherm models, but the Freundlich equation was more suitable. The adsorption was much easier at low temperature than that at high temperature at the initial Bt toxin concentration varying from 0 to 1 000 mg L~(-1). The negative values of the standard free energy (Δ_rG_m~θ) indicated that the adsorption of Bt toxin by minerals and soil colloids were spontaneous. The standard enthalpy changes (Δ_rH_m~θ) of the toxin adsorbed by montmorillinite was positive while that by others were negative, suggesting that the adsorption by montmorillinite was endothermic and by others were exothermic. The standard enthalpy changes (Δ_rH_m~θ) were all less than 40 kJ mol~(-1), showing that the adsorption was physicosorption.
2) The Bt toxin could be adsorbed easily by minerals and soil colloids, and the adsorption amount in the first 0.25 h was the 75%-80% of the equilibrium adsorption amount. The adsorption kinetic of Bt toxin can be described by the intra-particle diffusion model, however, the value of the contant C was not 0. This phenomenon showed that the speed of intra-particle diffusion was not the only dominant factor of the adsorption. Besides, the adsorption dynamic of Bt toxin consisted with the pseudo-first-order, pseudo-second-order, and Elovich equations in which the pseudo-second-order had the best fitting results, and it indicated that the adsorption is mainly controlled by the surface adsorption process.
3) The adsorption amount of the toxin were greatest at pH 6 by goethite, montmorillonite, kaolinite, yellow brown soil, and yellow cinnamon soil, while the maximum adsorption by silicon dioxide, red soil, and latosol presented at pH 7. The ]overall trend was the adsorption capacity of the toxin decreased with the pH increase, which is attributed to the isoelectric point (IEP) of Bt toxin and the point of zero charge (PZC) of minerals and soils.
4) The adsorption of Bt toxin by minerals and soil colloids were affected by low-molecular-weight organic acid ligands and inorganic salts. Low concentrations (< 10 mmol L~(-1)) of organic acid ligands (acetate, oxalate, citrate) inhibited toxin adsorption, whereas high concentrations promoted adsorption, and the inorganic salts had the opposite effects. The effect degree of the different organic acid ligands was oxalate > citrate > citrate, while H_2PO_4~- > NO_3~- and NH_4~+ > K~+ for inorganic ions.
5) The adsorbed Bt toxin was hardly desorbed. The desorption rate of the toxin by 0.1 mmol L~(-1) NaCl and phosphate was less than 5.3% and 13.1%, respectively, which indicated that a small proportion of the toxin adsorbed by minerals and soil colloids via electrostatic forces and ligand exchange. The desorption rates of Bt toxin were increased obviously when the organic acid ligands were present, the results suggested that organic acid ligands can markedly loose the bond of Bt toxin to minerals and soil colloids, and oxalate had the most significant effects among the tested organic acid ligands.
6) Two fluorescence peaks of Bt toxin presented at 338 ran of tryptophan residues and 314.5 nm of the tyrosine residues by the excitation at 282 nm. The tryptophan peak of the toxin desorbed from kaolinite, montmorillonite and soil colloids were red-shifted 5 to 9.5 nm, and their tyrosine peak remained at 314.5 nm without shift. However, the two peaks of the toxin desorbed from goethite and silicon dioxide didn't shift obviously. The red-shift of the fluorescence peaks suggested that the polarity of the microenvironment of the toxin increased during the adsorption and desorption. The CD spectra showed that theα-helix andβ-sheet content of the toxin after desorption from minerals and soil colloids increased while P-turn and random coil content decreased in comparison to native toxin, which indicated that the ordered structure (α-helix +β-sheet) of the toxin increased in a certain extent during the adsorption and desorption. The special IR spectra of amideⅠ(1653 cm~(-1)) and amideⅡ(1543 cm~(-1)) of the adsorbed toxin did not shifted obviously in comparision to the native toxin, and the peaks at 1055, 1240 and 2362 cm~(-1) were disappeared, these results indicated that the radicle of C-N played a key role in the adsorption of Bt toxin by minerals and soil colloids with no obvious changes in the secondary structure of Bt toxin after adsorption.
7) At 25℃, the remaining amount of the free Bt toxin (without soils) when the initial concentration was 0.3μg g~(-1) decreased as the cultivation time increase, and the remaining amount was less than 55% of the initial content after cultivating 10 days. The results indicated that the free toxin could degrade rapidly, and the degradation of the toxin with microorganisms (especially extracted from the soil planting non transgenic Bt plants) was more than that without microorganisms. However, as the cultivation time increase, the remaining of Bt toxin in soils (latosol, yellow brown soil and yellow cinnamon soil) increased firstly (1-2 days) and then gradually decreased. The half-life (DT_(50)) of Bt toxin in soil tested was about 10 days, and the effects of macroorganisms on the degradation of Bt toxin in soils were not obvious. The degradation rates of Bt toxin in yellow brown soil and yellow cinnamon soil were lower that that in latosol at cultivation time varying from 10 to 40 days, with no obvious differences of degradation rate in 3 tested soils at the first cultivation days (less than 10 days).
1)Bt毒素在供试胶体表面的等温吸附曲线为L型,可用Langmuir和Freundlich方程拟合,且Freundlich方程拟合效果更好。在Bt毒素初始浓度为0-1 000 mg L~(-1)时,其在矿物胶体表面的吸附量随温度升高而减少,在供试胶体表面吸附的标准自由能(Δ_rG_m~θ)为负值,表明该吸附是自发反应。Bt毒素除在蒙脱石表面吸附的标准焓变(Δ_rH_m~θ)为正值是吸热反应外,在其它供试胶体表面吸附的标准焓变均为负值,是放热反应;且所有吸附的标准焓变值均小于40 kJ mol~(-1),说明该吸附主要是物理吸附。
2)Bt毒素在供试胶体表面能快速吸附,0.25 h的吸附量约占平衡吸附量的64%-80%,1-2 h基本达平衡。该吸附可用粒子内扩散模型拟合,常数C值不为0,表明粒子内扩散速度不是Bt毒素在供试胶体表面吸附快慢的唯一控制因素。此外,其吸附动力学也符合一级方程、二级方程和Elovich方程,以二级动力学方程的拟合程度最好,说明该吸附主要受表面吸附控制。
3)针铁矿、蒙脱石、高岭石、黄棕壤和黄褐土等胶体在pH 6时对Bt毒素的吸附量最大,二氧化硅、红壤和砖红壤等胶体在pH 7时吸附量最大,其吸附量一般随pH的升高而降低,这主要与Bt毒素的等电点及供试胶体的电荷零点有关。
4)低分子量有机酸盐和无机配体对Bt毒素在供试胶体表面的吸附受的影响。对Bt毒素的吸附低浓度(<10 mmol L~(-1))有机酸盐(乙酸盐、草酸盐、柠檬酸盐)起抑制作用,高浓度有机酸盐起促进作用;不同有机酸盐抑制作用的强弱为草酸盐<柠檬酸盐≤乙酸盐。无机配体对Bt毒素吸附的影响与有机酸盐相反,低浓度无机盐促进其吸附,高浓度的起抑制作用,不同离子的影响为H_2PO_4~->NO_3~-,NH_4~+>K~+。
5)吸附态Bt毒素很难被解吸。用0.1 mmol L~(-1) NaCl作解吸剂时,供试胶体表面吸附态Bt毒素的解吸率<5.3%,磷酸缓冲液作解吸剂时其解吸率<13.1%,表明它只有少部分以静电和配位的方式吸附在供试胶体表面。在有低分子量有机酸盐等配体存在条件下,NaCl和磷酸缓冲液对吸附态Bt毒素的解吸率均有显著增加(除乙酸盐体系中蒙脱石和砖红壤吸附态Bt毒素外),表明有机酸盐的存在一定程度上增加了静电和配位吸附,影响大小为草酸盐>柠檬酸盐>乙酸盐。
6)荧光光谱显示Bt毒素在282 nm激发光的作用下有酪氨酸(314.5 nm)和色氨酸(338 nm)荧光峰,其与高岭石、蒙脱石及土壤等胶体作用后色氨酸荧光峰红移了5-9.5 nm,酪氨酸荧光峰没有变化;与针铁矿和二氧化硅胶体作用后色氨酸和酪氨酸的荧光峰蓝移了0.5-1 nm,变化不明显。荧光峰的红移表明Bt毒素的吸附导致其内源荧光基团所处的分子环境极性增强。圆二色光谱表明吸附导致Bt毒素的二级结构改变,α螺旋和β折叠含量增加,而β转角和无规则卷曲含量降低。红外光谱显示,吸附后Bt毒素的酰胺Ⅰ带(1653 cm~(-1))和酰胺Ⅱ带(1543 cm~(-1))等特征光谱峰未发生明显位移,只是1055,1240和2362 cm~(-1)处的峰消失,表明C-N残基在供试胶体吸附Bt毒素中起重要作用,且吸附后毒素的二级结构变化不明显。
7)25℃条件下,初始浓度为0.3μg g~(-1)的游离态Bt毒素随时间增加残留量逐渐减少,10天时残留量小于原始加入量的55%,培养40天后约降解了80%,表明游离态Bt毒素降解速率较快。有外源微生物存在,其降解速率更快,表明微生物能利用游离态Bt毒素;双指数模型拟合结果表明,微生物存在下其降解速率更快,且从种植非转Bt植物上壤中提取的微生物对游离态Bt毒素的利用更快。Bt毒素在供试灭菌土壤(砖红壤、黄褐土和黄棕壤)中的残留量随着培养时间的推移先增加(1-2天)后逐步减少,降解50%的时间(DT_(50))为10天左右,外源微生物对土壤中Bt毒素的降解速率影响不明显。初始阶段Bt毒素在3种土壤中降解速度的差异不明显,而后期在黄棕壤和黄褐土中的降解速率比在砖红壤中的慢。
The wide-spread cultivation of transgenic plants would pose a risk to natural and agricultural ecosystems, especially the transgenic Bacillus thuringinesis (Bt) plants, for they release Bt toxin to the soil ecosystem by root exudates, plant biomass and pollen. This study investigated the adsorption and desorption of Bt toxin by montmorillinite, kaolinite, goethite, silicon dioxide, red soil (Ultisol), latosol (Oxisol), yellow brown soil (Alfisol) and yellow cinnamon soil (Alfisol). The kinetic and thermodynamic parameters were analyzed to understand the adsorption mechanisms, while infrared spectra, circular dichroism (CD) spectra and fluorescence spectra were used to investigate the secondary structure changes of Bt toxin during the adsorption and desorption process. The remaining of Bt toxin in soils with or without microorganisms were also studied. This investigation helps to evaluate the behavior and fate of Bt toxins in the soil ecosystem. The main results are described below.
1) The isotherm adsorption curves were L-type, and the adsorption data fitted well with both Langmuir and Freundlich isotherm models, but the Freundlich equation was more suitable. The adsorption was much easier at low temperature than that at high temperature at the initial Bt toxin concentration varying from 0 to 1 000 mg L~(-1). The negative values of the standard free energy (Δ_rG_m~θ) indicated that the adsorption of Bt toxin by minerals and soil colloids were spontaneous. The standard enthalpy changes (Δ_rH_m~θ) of the toxin adsorbed by montmorillinite was positive while that by others were negative, suggesting that the adsorption by montmorillinite was endothermic and by others were exothermic. The standard enthalpy changes (Δ_rH_m~θ) were all less than 40 kJ mol~(-1), showing that the adsorption was physicosorption.
2) The Bt toxin could be adsorbed easily by minerals and soil colloids, and the adsorption amount in the first 0.25 h was the 75%-80% of the equilibrium adsorption amount. The adsorption kinetic of Bt toxin can be described by the intra-particle diffusion model, however, the value of the contant C was not 0. This phenomenon showed that the speed of intra-particle diffusion was not the only dominant factor of the adsorption. Besides, the adsorption dynamic of Bt toxin consisted with the pseudo-first-order, pseudo-second-order, and Elovich equations in which the pseudo-second-order had the best fitting results, and it indicated that the adsorption is mainly controlled by the surface adsorption process.
3) The adsorption amount of the toxin were greatest at pH 6 by goethite, montmorillonite, kaolinite, yellow brown soil, and yellow cinnamon soil, while the maximum adsorption by silicon dioxide, red soil, and latosol presented at pH 7. The ]overall trend was the adsorption capacity of the toxin decreased with the pH increase, which is attributed to the isoelectric point (IEP) of Bt toxin and the point of zero charge (PZC) of minerals and soils.
4) The adsorption of Bt toxin by minerals and soil colloids were affected by low-molecular-weight organic acid ligands and inorganic salts. Low concentrations (< 10 mmol L~(-1)) of organic acid ligands (acetate, oxalate, citrate) inhibited toxin adsorption, whereas high concentrations promoted adsorption, and the inorganic salts had the opposite effects. The effect degree of the different organic acid ligands was oxalate > citrate > citrate, while H_2PO_4~- > NO_3~- and NH_4~+ > K~+ for inorganic ions.
5) The adsorbed Bt toxin was hardly desorbed. The desorption rate of the toxin by 0.1 mmol L~(-1) NaCl and phosphate was less than 5.3% and 13.1%, respectively, which indicated that a small proportion of the toxin adsorbed by minerals and soil colloids via electrostatic forces and ligand exchange. The desorption rates of Bt toxin were increased obviously when the organic acid ligands were present, the results suggested that organic acid ligands can markedly loose the bond of Bt toxin to minerals and soil colloids, and oxalate had the most significant effects among the tested organic acid ligands.
6) Two fluorescence peaks of Bt toxin presented at 338 ran of tryptophan residues and 314.5 nm of the tyrosine residues by the excitation at 282 nm. The tryptophan peak of the toxin desorbed from kaolinite, montmorillonite and soil colloids were red-shifted 5 to 9.5 nm, and their tyrosine peak remained at 314.5 nm without shift. However, the two peaks of the toxin desorbed from goethite and silicon dioxide didn't shift obviously. The red-shift of the fluorescence peaks suggested that the polarity of the microenvironment of the toxin increased during the adsorption and desorption. The CD spectra showed that theα-helix andβ-sheet content of the toxin after desorption from minerals and soil colloids increased while P-turn and random coil content decreased in comparison to native toxin, which indicated that the ordered structure (α-helix +β-sheet) of the toxin increased in a certain extent during the adsorption and desorption. The special IR spectra of amideⅠ(1653 cm~(-1)) and amideⅡ(1543 cm~(-1)) of the adsorbed toxin did not shifted obviously in comparision to the native toxin, and the peaks at 1055, 1240 and 2362 cm~(-1) were disappeared, these results indicated that the radicle of C-N played a key role in the adsorption of Bt toxin by minerals and soil colloids with no obvious changes in the secondary structure of Bt toxin after adsorption.
7) At 25℃, the remaining amount of the free Bt toxin (without soils) when the initial concentration was 0.3μg g~(-1) decreased as the cultivation time increase, and the remaining amount was less than 55% of the initial content after cultivating 10 days. The results indicated that the free toxin could degrade rapidly, and the degradation of the toxin with microorganisms (especially extracted from the soil planting non transgenic Bt plants) was more than that without microorganisms. However, as the cultivation time increase, the remaining of Bt toxin in soils (latosol, yellow brown soil and yellow cinnamon soil) increased firstly (1-2 days) and then gradually decreased. The half-life (DT_(50)) of Bt toxin in soil tested was about 10 days, and the effects of macroorganisms on the degradation of Bt toxin in soils were not obvious. The degradation rates of Bt toxin in yellow brown soil and yellow cinnamon soil were lower that that in latosol at cultivation time varying from 10 to 40 days, with no obvious differences of degradation rate in 3 tested soils at the first cultivation days (less than 10 days).
引文
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