土壤腐殖酸与酶蛋白相互作用的机制
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摘要
蛋白质是环境中重要的活性组分,主要通过动植物和微生物的分泌、残体遗骸的分解释放、人类的活动等进入土壤和水体环境中。这些蛋白质既包含具有催化作用的脲酶、酸性磷酸酶等,也包含一些对环境存在潜在危害性的朊病毒、禽流感病毒等。当它们进入环境中,很少以游离态存在,常与有机物、矿物、微生物等结合在一起。腐殖酸作为环境中重要的组分,广泛存在于土壤、水体等环境中,可以在静电、疏水、化学键等的驱使下以吸附、共聚或者包埋等作用机制与蛋白质发生作用。本文主要以腐殖酸和溶菌酶为研究对象,从胶体性质的角度出发,在了解两者表面电荷性质的基础上,通过研究pH值、离子强度、复合物质量比、腐殖酸类型等对两者作用过程中电化学性质以及对酶活性、结构的影响,进一步揭示两者的主要作用机制。主要结果如下：
1.阐明了胶体表面绝对电荷量的测定方法,在酸碱滴定和Stat-pH滴定的基础上,通过测定曲线上某一点处的绝对电荷量,经过平移最终得到了不同离子强度下腐殖酸表面绝对电荷量随pH值变化的曲线。绝对电荷的测定方法需要根据样品的性质来选择,本文对采用聚电解质滴定法和采用改变样品溶液pH值和离子强度计算质子解离量来测定腐殖酸表面绝对电荷量的两种方法进行了比较,结果发现对于胡敏酸而言,聚电解质滴定法测得的结果大于酸碱滴定的结果；富里酸不适合采用聚电解质滴定法进行测定,测定结果偏低。
2.采用颗粒电荷滴定仪测定了腐殖酸滴定溶菌酶的过程中电势(mV)以及pH值的变化。根据电势的变化计算达到复合物等电点时腐殖酸与溶菌酶的电中性质量比(mHS/mLSZ)IEp,结果表明腐殖酸与溶菌酶之间具有较高的分子亲和力,电中性质量比(mHS/mLSZ)IEP随离子强度的增加而增加,随pH值的升高而下降。并与理论得到的电荷补偿质量比(mHS/mLSZ)CCP进行了比较,结果发现(mHS/mLSZ) IEP>(mHS/mLSZ) ccp,这是因为K+参与复合物的形成,且其参与量随着离子强度的增加而增加,随着pH值的升高而下降。此外,K+的参与量还与腐殖酸的类型有关,在JGFA与溶菌酶复合物中的参与量最大,而在PAHA与溶菌酶复合物中的参与量最小。
3.观察了不同质量比的腐殖酸与溶菌酶复合物随时间变化的絮凝现象,结果表明腐殖酸与溶菌酶的复合物在等电点处的絮凝现象最明显,较短的时间内出现了大量的絮凝物。对于胡敏酸与溶菌酶的复合物而言,较大的质量比不易于絮凝物的产生,带负电荷的复合物分子之间存在的静电排斥力促进了复合物解聚的发生,抑制了絮凝物的产生。
4.腐殖酸与蛋白质的作用过程中,通过测定复合物中酶活性的变化来反映溶液pH值、离子强度、复合质量比对复合物形成、电荷性质、结构等方面的影响,从而进一步揭示两者的作用机制。结果表明静电作用力在腐殖酸与蛋白质之间起着重要的作用。在静电引力的驱动下,溶菌酶被腐殖酸包被在其分子结构中,底物接触活性位点受阻导致活性降低。在两者的作用过程中,复合物的质量比起着重要的作用,决定了复合物表面所带的电荷量以及复合物分子的结构。此外,离子强度对腐殖酸和溶菌酶之间的静电屏蔽作用与质量比有关,在等电点前后其对酶活的作用相反,进一步表明静电作用在两者之间的重要性。富里酸分子量低、结构简单,对溶菌酶包被的程度以及对酶活性的抑制程度均小于胡敏酸。腐殖酸与两性分子脲酶的静电作用受溶液pH值的影响：pH值大于脲酶等电点时,两者之间存在静电排斥作用；pH值小于脲酶等电点时,两者之间主要以静电引力为主。离子强度对腐殖酸和脲酶之间的静电作用力同样具有屏蔽作用。腐殖酸降低了溶菌酶的稳定性,且在电中性质量比处稳定性最低。相反,腐殖酸的加入减弱了脲酶分子之间的碰撞从而提高了脲酶的稳定性。
5.采用荧光光谱法研究了腐殖酸与溶菌酶的作用对色氨酸微环境的改变,结果发现pH5及5mmol L-1KCl时,腐殖酸对溶菌酶在340nm处的荧光峰产生猝灭作用(激发波长为280nm)。随着复合物质量比的增加,荧光猝灭的程度增强,当质量比大于0.2时,溶菌酶在该处的荧光发射峰几乎消失；340nm处的荧光峰主要由色氨酸贡献,随着腐殖酸质量的增加,色氨酸的荧光发射峰逐渐发生红移,表明腐殖酸与溶菌酶的结合使得色氨酸残基附近的微环境发生了变化,极性增强,疏水性降低,蛋白质的结构变得疏松。
6.采用圆二色谱法测定了在腐殖酸与溶菌酶复合过程中,蛋白质二级结构的变化,结果发现PAHA、JGHA与溶菌酶的复合物中,α螺旋、自由卷曲的含量均随着质量比的增加呈先下降后上升的趋势,在等电点处形成一个拐点,与复合物酶活性随质量比的变化趋势一致。β折叠的含量随质量比的增加呈先上升后下降的变化趋势,同样在等电点处形成一个拐点,拐点为胡敏酸对溶菌酶包被和解聚的一个转折点。对于JGFA来说,α螺旋、转角、自由卷曲的含量均随着复合物质量比的增加而增加,而β折叠则随着质量比的增加呈现下降的趋势,在电中性质量比f=0.6时均未出现拐点。
7.采用衰减全反射红外光谱法研究了复合时间、质量比、pH值、离子强度等对复合物红外光谱的影响。当质量比f=(mHA/mLSZ)IEP时,腐殖酸对酶的包被程度达到最大,溶菌酶的红外吸收峰均受到了较大程度的抑制。随着pH值的升高,胡敏酸对溶菌酶的红外吸收峰产生最大屏蔽作用时对应的质量比降低,与复合物电中性质量比随pH值的升高而下降是一致的。当离子强度增加至50mmol L-1时,胡敏酸对溶菌酶的包被及解聚现象不明显,不同质量比复合物的红外光谱差异不大。质量比、pH值、离子强度对富里酸与溶菌酶复合物红外光谱的影响都不大。
Protein as a component of organic matter, can enter into the soil and water system through the secretion of animals, plants roots and microorganisms, the release from the residue and decomposition of these living things, and human's activities. Enzymes, e.g. phosphatase and urease, play important roles in the environment, especially for the geochemical cycle of elements such as phosphorus (P), nitrogen (N) etc. Another type of protein like prion causing transmissible spongiform encephalopathies, bacillus thuringiensis (Bt) protein, avian influenza virus, are potentially harmful factors in environment for human and animals. When these proteins are released into environment, they will combine with organic matter, minerals and microorganisms. Humic Substances (HS) are important components of organic matter and are widely distributed in the soil and water. Driven by the electrostatic interaction, hydrophobic force and chemical binding, proteins can be adsorbed onto or entrapped into HS, or can co-polymerize with HS.In this paper, we firstly investigated the colloidal properties of HS and lysozyme (LSZ), especially the surface charge of them. Then we studied the effect of HS on the activity, stability and structure of LSZ. Factors like pH, ionic strength, mass ratio HS/LSZ and types of humic acids were considered. Main results are below:
1. We elucidated the method of getting the abosulte charge of colloids. On the basement of acid-base titration and Stat-pH titration, we tested the absolute charge of one point on△charge-pH curve and then moved them parallel using this point as reference. Finally we can obtain absolute charge of colloids as a function of pH at different KC1concentrations. For getting the absolute charge of HS, two methods were applied and compared. The first method was polyDADMAC titration and the absolute charge of HS was calculated according to the mass ratio polyDADMAC/HS at IEP of the complex and the charge of polyDADMAC (5.9mmol g-1); the second method was calculating the proton release by changing pH and ionic strength of protonated sample solutions. The results showed that the charge of JGHA obtained from the first method was larger than the second and for JGFA the first method was not suitable and made the result much lower.
2. We monitored the change of potential (mV) and pH values when LSZ was titrated by HS. The mass ratio HS/LSZ at IEP of complex (mHS/mLSZ)IEP can be obtained with muitek particle charge detector. The results showed that the affinity of HS and LSZ was high and (mHS/mLSZ)IEP increased with increase of ionic strength and decrease of pH values. Moreover, we also calculated the mass ratio (mHS/mLSZ)CCP, which equals the charge density of LSZ divided by the charge density of HS. By comparison,(mHS/mLSZ)IEP>(mHS/mLSZ)CCP can be found, which is due to the participation of K+And the participation of K+increased with ionic strength and decreased with pH. Moreover, it was also related to the types of HS. The participation of K+reached the largest in JGFA-LSZ complexes and the lowest in PAHA-LSZ complexes.
3. The flocculation of HS-LSZ complexes (with series of mass ratios) as a function of time was observed. The results showed that the aggregation of complex at IEP occurred the earliest and the amount of aggregation was the largest. For HA-LSZ complexes with larger mass ratio (f=1.0), the aggregation was inhibited. This was probably due to the electrostatic repulsion between negatively charged molecules.
4. During HS interaction with LSZ, the enzyme activity in complexes was measured to reflect the effect of pH, ionic strength, mass ratio HS/enzyme on the formation, charge properties and structure of complexes, further indicating the mechanism of their interaction. Our results showed that electrostatic interaction plays an important role in the interaction of HS with LSZ. Driven by electrostatic attraction, LSZ was encapsulated into HS inner structure. As a result, the active sites of LSZ were shielded and substrate could not contact with them, leading the decrease of LSZ activity. Especially for two oppsitely charged particles, mass ratio was crucial because it determined the surface charge and structure of complexes; ionic strength screened the electrostatic interaction between HS and LSZ, which was also related to mass ratio. The effect of ionic strength on LSZ activity was opposite before and after IEP of the complex, indicating the importance of electrostatic interaction. Due to the smaller molecular weight and simpler structure of FA, the encapsulation of LSZ by FA was much weaker. The interaction of HS with amphoteric urease was also investigated. The electrostatic interaction was also important and was dependent on pH value of solution. When pH was larger than the PZC of urease, electrostatic repulsion hindered the encapsulation of urease by HS and maintained the high activity of urease; when pH was lower than the PZC of urease, electrostatic attraction improve the interaction of HS with urease, which was similar with LSZ-HS complex. The presence of HS decreased LSZ stability, and this inhibition reached maximum at the IEP of complexes due to the strongest aggregation of complexes at this point. However, the presence of HS improved the stability of urease. This was possibly because the collision among urease moleculars was decreased by HS and increased its stability.
5. The fluorescence spectrum and synchronous excitation fluorescence spectrum (△λ,=60) of LSZ in the presence of HS at pH5and5mmol L-1KC1were conducted in order to investigate the change of microenvironment of LSZ. The results indicated that with excitation wave set at280nm, a progressive reduction in fluorescence intensity at340nm was caused by the increasing concentrations of HS. As mass ratio increased to0.2, the fluorescence intensity was nearly zero. Thus, the fluorescence of LSZ was strongly quenched and the red shift of λmax can be reasonably attributed to an increased polarity (or a decreased hydrophobicity) and loose structure of the region surrounding Trp site.
6. The circular dichroism spectra of LSZ in the presence of HS were conducted and the secondary structure of LSZ in HS-LSZ complexes was calculated. Our results showed that the content of a-helix and random coils of HA-LSZ complexes firstly increased and then decreased with the increase of mass ratio HS/LSZ. For β-sheets, its content as a function of mass ratio HS/LSZ was opposite to a-helix and random coils. The turning points for the percentage of a-helix, random coils and β-sheets as a function of mass ratio HS/LSZ were formed at IEP of the complex, which was corresponding with the acticity of LSZ in complex as a function of mass ratio HS/LSZ. At this point, the complex of HS-LSZ started to disaggregate, accompanied with the change of secondard structure. For JGFA-LSZ complex, the content of a-helix, random coils and β-turns increased with increasing mass ratio while the content of β-sheets decreased.
7. With ATR-FTIT spectrum, we investigated the effect of HS on the FTIR spectrum of LSZ at different conditions like incubation time, pH, ionic strength and mass ratio HS/LSZ. The results showed that the adsorption bands of LSZ were screened by HS most significantly around the IEP of HS-LSZ complex, Moreover, the mass ratio at which the screening was the most significant decreased with the increase of pH. This was corresponding to the decrease of (HS/LSZ)IEP with increasing pH. When ionic strength was increased to50mmol L-1, the similar results were not observed. As to JGFA-LSZ complex, factors like mass ratio, pH and ionic strength had little influence.
1.阐明了胶体表面绝对电荷量的测定方法,在酸碱滴定和Stat-pH滴定的基础上,通过测定曲线上某一点处的绝对电荷量,经过平移最终得到了不同离子强度下腐殖酸表面绝对电荷量随pH值变化的曲线。绝对电荷的测定方法需要根据样品的性质来选择,本文对采用聚电解质滴定法和采用改变样品溶液pH值和离子强度计算质子解离量来测定腐殖酸表面绝对电荷量的两种方法进行了比较,结果发现对于胡敏酸而言,聚电解质滴定法测得的结果大于酸碱滴定的结果；富里酸不适合采用聚电解质滴定法进行测定,测定结果偏低。
2.采用颗粒电荷滴定仪测定了腐殖酸滴定溶菌酶的过程中电势(mV)以及pH值的变化。根据电势的变化计算达到复合物等电点时腐殖酸与溶菌酶的电中性质量比(mHS/mLSZ)IEp,结果表明腐殖酸与溶菌酶之间具有较高的分子亲和力,电中性质量比(mHS/mLSZ)IEP随离子强度的增加而增加,随pH值的升高而下降。并与理论得到的电荷补偿质量比(mHS/mLSZ)CCP进行了比较,结果发现(mHS/mLSZ) IEP>(mHS/mLSZ) ccp,这是因为K+参与复合物的形成,且其参与量随着离子强度的增加而增加,随着pH值的升高而下降。此外,K+的参与量还与腐殖酸的类型有关,在JGFA与溶菌酶复合物中的参与量最大,而在PAHA与溶菌酶复合物中的参与量最小。
3.观察了不同质量比的腐殖酸与溶菌酶复合物随时间变化的絮凝现象,结果表明腐殖酸与溶菌酶的复合物在等电点处的絮凝现象最明显,较短的时间内出现了大量的絮凝物。对于胡敏酸与溶菌酶的复合物而言,较大的质量比不易于絮凝物的产生,带负电荷的复合物分子之间存在的静电排斥力促进了复合物解聚的发生,抑制了絮凝物的产生。
4.腐殖酸与蛋白质的作用过程中,通过测定复合物中酶活性的变化来反映溶液pH值、离子强度、复合质量比对复合物形成、电荷性质、结构等方面的影响,从而进一步揭示两者的作用机制。结果表明静电作用力在腐殖酸与蛋白质之间起着重要的作用。在静电引力的驱动下,溶菌酶被腐殖酸包被在其分子结构中,底物接触活性位点受阻导致活性降低。在两者的作用过程中,复合物的质量比起着重要的作用,决定了复合物表面所带的电荷量以及复合物分子的结构。此外,离子强度对腐殖酸和溶菌酶之间的静电屏蔽作用与质量比有关,在等电点前后其对酶活的作用相反,进一步表明静电作用在两者之间的重要性。富里酸分子量低、结构简单,对溶菌酶包被的程度以及对酶活性的抑制程度均小于胡敏酸。腐殖酸与两性分子脲酶的静电作用受溶液pH值的影响：pH值大于脲酶等电点时,两者之间存在静电排斥作用；pH值小于脲酶等电点时,两者之间主要以静电引力为主。离子强度对腐殖酸和脲酶之间的静电作用力同样具有屏蔽作用。腐殖酸降低了溶菌酶的稳定性,且在电中性质量比处稳定性最低。相反,腐殖酸的加入减弱了脲酶分子之间的碰撞从而提高了脲酶的稳定性。
5.采用荧光光谱法研究了腐殖酸与溶菌酶的作用对色氨酸微环境的改变,结果发现pH5及5mmol L-1KCl时,腐殖酸对溶菌酶在340nm处的荧光峰产生猝灭作用(激发波长为280nm)。随着复合物质量比的增加,荧光猝灭的程度增强,当质量比大于0.2时,溶菌酶在该处的荧光发射峰几乎消失；340nm处的荧光峰主要由色氨酸贡献,随着腐殖酸质量的增加,色氨酸的荧光发射峰逐渐发生红移,表明腐殖酸与溶菌酶的结合使得色氨酸残基附近的微环境发生了变化,极性增强,疏水性降低,蛋白质的结构变得疏松。
6.采用圆二色谱法测定了在腐殖酸与溶菌酶复合过程中,蛋白质二级结构的变化,结果发现PAHA、JGHA与溶菌酶的复合物中,α螺旋、自由卷曲的含量均随着质量比的增加呈先下降后上升的趋势,在等电点处形成一个拐点,与复合物酶活性随质量比的变化趋势一致。β折叠的含量随质量比的增加呈先上升后下降的变化趋势,同样在等电点处形成一个拐点,拐点为胡敏酸对溶菌酶包被和解聚的一个转折点。对于JGFA来说,α螺旋、转角、自由卷曲的含量均随着复合物质量比的增加而增加,而β折叠则随着质量比的增加呈现下降的趋势,在电中性质量比f=0.6时均未出现拐点。
7.采用衰减全反射红外光谱法研究了复合时间、质量比、pH值、离子强度等对复合物红外光谱的影响。当质量比f=(mHA/mLSZ)IEP时,腐殖酸对酶的包被程度达到最大,溶菌酶的红外吸收峰均受到了较大程度的抑制。随着pH值的升高,胡敏酸对溶菌酶的红外吸收峰产生最大屏蔽作用时对应的质量比降低,与复合物电中性质量比随pH值的升高而下降是一致的。当离子强度增加至50mmol L-1时,胡敏酸对溶菌酶的包被及解聚现象不明显,不同质量比复合物的红外光谱差异不大。质量比、pH值、离子强度对富里酸与溶菌酶复合物红外光谱的影响都不大。
Protein as a component of organic matter, can enter into the soil and water system through the secretion of animals, plants roots and microorganisms, the release from the residue and decomposition of these living things, and human's activities. Enzymes, e.g. phosphatase and urease, play important roles in the environment, especially for the geochemical cycle of elements such as phosphorus (P), nitrogen (N) etc. Another type of protein like prion causing transmissible spongiform encephalopathies, bacillus thuringiensis (Bt) protein, avian influenza virus, are potentially harmful factors in environment for human and animals. When these proteins are released into environment, they will combine with organic matter, minerals and microorganisms. Humic Substances (HS) are important components of organic matter and are widely distributed in the soil and water. Driven by the electrostatic interaction, hydrophobic force and chemical binding, proteins can be adsorbed onto or entrapped into HS, or can co-polymerize with HS.In this paper, we firstly investigated the colloidal properties of HS and lysozyme (LSZ), especially the surface charge of them. Then we studied the effect of HS on the activity, stability and structure of LSZ. Factors like pH, ionic strength, mass ratio HS/LSZ and types of humic acids were considered. Main results are below:
1. We elucidated the method of getting the abosulte charge of colloids. On the basement of acid-base titration and Stat-pH titration, we tested the absolute charge of one point on△charge-pH curve and then moved them parallel using this point as reference. Finally we can obtain absolute charge of colloids as a function of pH at different KC1concentrations. For getting the absolute charge of HS, two methods were applied and compared. The first method was polyDADMAC titration and the absolute charge of HS was calculated according to the mass ratio polyDADMAC/HS at IEP of the complex and the charge of polyDADMAC (5.9mmol g-1); the second method was calculating the proton release by changing pH and ionic strength of protonated sample solutions. The results showed that the charge of JGHA obtained from the first method was larger than the second and for JGFA the first method was not suitable and made the result much lower.
2. We monitored the change of potential (mV) and pH values when LSZ was titrated by HS. The mass ratio HS/LSZ at IEP of complex (mHS/mLSZ)IEP can be obtained with muitek particle charge detector. The results showed that the affinity of HS and LSZ was high and (mHS/mLSZ)IEP increased with increase of ionic strength and decrease of pH values. Moreover, we also calculated the mass ratio (mHS/mLSZ)CCP, which equals the charge density of LSZ divided by the charge density of HS. By comparison,(mHS/mLSZ)IEP>(mHS/mLSZ)CCP can be found, which is due to the participation of K+And the participation of K+increased with ionic strength and decreased with pH. Moreover, it was also related to the types of HS. The participation of K+reached the largest in JGFA-LSZ complexes and the lowest in PAHA-LSZ complexes.
3. The flocculation of HS-LSZ complexes (with series of mass ratios) as a function of time was observed. The results showed that the aggregation of complex at IEP occurred the earliest and the amount of aggregation was the largest. For HA-LSZ complexes with larger mass ratio (f=1.0), the aggregation was inhibited. This was probably due to the electrostatic repulsion between negatively charged molecules.
4. During HS interaction with LSZ, the enzyme activity in complexes was measured to reflect the effect of pH, ionic strength, mass ratio HS/enzyme on the formation, charge properties and structure of complexes, further indicating the mechanism of their interaction. Our results showed that electrostatic interaction plays an important role in the interaction of HS with LSZ. Driven by electrostatic attraction, LSZ was encapsulated into HS inner structure. As a result, the active sites of LSZ were shielded and substrate could not contact with them, leading the decrease of LSZ activity. Especially for two oppsitely charged particles, mass ratio was crucial because it determined the surface charge and structure of complexes; ionic strength screened the electrostatic interaction between HS and LSZ, which was also related to mass ratio. The effect of ionic strength on LSZ activity was opposite before and after IEP of the complex, indicating the importance of electrostatic interaction. Due to the smaller molecular weight and simpler structure of FA, the encapsulation of LSZ by FA was much weaker. The interaction of HS with amphoteric urease was also investigated. The electrostatic interaction was also important and was dependent on pH value of solution. When pH was larger than the PZC of urease, electrostatic repulsion hindered the encapsulation of urease by HS and maintained the high activity of urease; when pH was lower than the PZC of urease, electrostatic attraction improve the interaction of HS with urease, which was similar with LSZ-HS complex. The presence of HS decreased LSZ stability, and this inhibition reached maximum at the IEP of complexes due to the strongest aggregation of complexes at this point. However, the presence of HS improved the stability of urease. This was possibly because the collision among urease moleculars was decreased by HS and increased its stability.
5. The fluorescence spectrum and synchronous excitation fluorescence spectrum (△λ,=60) of LSZ in the presence of HS at pH5and5mmol L-1KC1were conducted in order to investigate the change of microenvironment of LSZ. The results indicated that with excitation wave set at280nm, a progressive reduction in fluorescence intensity at340nm was caused by the increasing concentrations of HS. As mass ratio increased to0.2, the fluorescence intensity was nearly zero. Thus, the fluorescence of LSZ was strongly quenched and the red shift of λmax can be reasonably attributed to an increased polarity (or a decreased hydrophobicity) and loose structure of the region surrounding Trp site.
6. The circular dichroism spectra of LSZ in the presence of HS were conducted and the secondary structure of LSZ in HS-LSZ complexes was calculated. Our results showed that the content of a-helix and random coils of HA-LSZ complexes firstly increased and then decreased with the increase of mass ratio HS/LSZ. For β-sheets, its content as a function of mass ratio HS/LSZ was opposite to a-helix and random coils. The turning points for the percentage of a-helix, random coils and β-sheets as a function of mass ratio HS/LSZ were formed at IEP of the complex, which was corresponding with the acticity of LSZ in complex as a function of mass ratio HS/LSZ. At this point, the complex of HS-LSZ started to disaggregate, accompanied with the change of secondard structure. For JGFA-LSZ complex, the content of a-helix, random coils and β-turns increased with increasing mass ratio while the content of β-sheets decreased.
7. With ATR-FTIT spectrum, we investigated the effect of HS on the FTIR spectrum of LSZ at different conditions like incubation time, pH, ionic strength and mass ratio HS/LSZ. The results showed that the adsorption bands of LSZ were screened by HS most significantly around the IEP of HS-LSZ complex, Moreover, the mass ratio at which the screening was the most significant decreased with the increase of pH. This was corresponding to the decrease of (HS/LSZ)IEP with increasing pH. When ionic strength was increased to50mmol L-1, the similar results were not observed. As to JGFA-LSZ complex, factors like mass ratio, pH and ionic strength had little influence.
引文
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