真菌漆酶高产菌株的发酵产酶及酶促降解有机染料的动力学研究
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摘要
白腐菌是对木质素降解能力最强的木腐真菌，是目前已知的能在一定条件下将木质素彻底降解为CO2和H2O的唯一一类微生物。白腐菌分泌的木素降解酶主要包括木质素过氧化物酶、锰过氧化物酶和漆酶。其中漆酶是一种含铜的多酚氧化酶，具有非常广泛的底物范围，可以催化氧化酚类和芳胺类化合物脱去羟基上的电子或质子，形成自由基，导致酚类及木素类化合物降解，同时分子氧被还原为水。在某些小分子化合物作为介体存在的条件下，漆酶还能够氧化非酚型木质素结构。近几年，随着研究的不断深入，白腐菌漆酶对木质素和与木质素结构相似的许多环境污染物的降解作用越来越受到科研工作者的关注，特别是在纸浆生物漂白、工业废水处理、有机染料脱色和高分子催化合成等方面，表现出了很大的研究价值和应用潜力。但是国内外至今还尚未有漆酶规模化生产的研究报道，漆酶的产量还远远不能满足上述工业应用的需要，并且价格比较昂贵。因此，目前需要解决的一个关键性问题是如何在控制成本的前提下提高漆酶的产量。
白腐菌Panus conchatus（贝壳状革耳菌）是一种常见的可食用和药用的野生真菌，对木质素的降解具有较强的选择性。本研究将白腐菌P. conchatus漆酶的生产由实验室摇瓶培养扩大到7.5L机械搅拌式发酵罐中，并对搅拌转速、通气量、温度等影响液体深层发酵的关键因素进行了考察和优化。结果表明，麦麸和硫酸铜能够显著提高摇瓶发酵中白腐菌P. conchatus漆酶的产量，缩短发酵周期，发酵液中漆酶活力最高可达196.1U/mL。通气量、培养温度和搅拌转速对白腐菌P. conchatus在机械搅拌式发酵罐中的产酶效率影响较大，最佳发酵条件为通气量1.0vvm，温度30°C，转速300rpm，漆酶活力在发酵22d时达到最大值约200U/mL，保持了和摇瓶发酵相同的产酶水平。Logistic模型能够较为准确的描述和预测白腐菌P. conchatus产漆酶的动力学过程，酶活测定值与方程计算值的线性相关系数R2达到了0.95以上。将该漆酶用于芦苇浆的生物漂白，纸浆经漆酶/介体系统处理后，卡伯值降低，可漂性有所提高，有利于后续漂白工段对纸浆中残余木素的脱除，提高漂白浆的白度。
以农业加工副产物和蔗渣酸法蒸煮废液为基础，研究用于白腐菌培养和漆酶生产的廉价原材料。结果表明，豆粕最适宜作为发酵底物进行漆酶的生产，液体培养基中漆酶最高活力达到476.7U/mL。木糖是蔗渣酸法蒸煮废水中最主要的单糖类物质，白腐菌Panus conchatus，Flammulina velutipes和Psathyrella candolleana都可在未经脱毒处理的废水制备的产酶培养基中生长，其中F. velutipes对废水毒性的抵抗力较强，其生物量和最高漆酶活力分别达到26.3g/L和200U/L。
通过盐析、超滤、离子交换色谱和凝胶过滤色谱等方法，将白腐菌P. conchatus漆酶纯化至电泳纯级别，其比活力达到912.3U/mg，比原始粗酶液中漆酶比活力提高了6.77倍，总酶活得率为74.1%。该漆酶的分子量约为65kDa，纯化后漆酶显蓝色，其紫外可见光谱学特征表明该漆酶为典型的真菌漆酶。漆酶对底物ABTS的米氏常数Km为5.7μ2，最大反应速度Vmax为31.06mM/(QMR)，最适反应温度和pH值分别为60oC和2.5，漆酶活性在4oC，pH8.0的环境中具有较好的稳定性，保存26d后，漆酶活力还能够保持为初始酶活的约97.9%。白腐菌P. conchatus漆酶可直接对蒽醌染料活性亮蓝RBBR和偶氮染料甲基橙进行脱色，粗酶液处理RBBR0.5h后，染料脱色率达到95.2%；处理甲基橙2h后，脱色率为84.1%。在介体HOBT存在的条件下，三苯甲烷类染料酸性品红可被粗酶液在1h内完全脱色。
白腐菌P. conchatus漆酶可有效降解蒽醌染料RBBR，反应条件为温度30°C，pH值4.0，RBBR初始浓度100mg/L，漆酶用量1U/mL时，反应15min后RBBR降解率即达到90%以上。根据降解过程中RBBR降解率与时间、温度、染料浓度以及漆酶用量等的关系，建立了白腐菌P. conchatus漆酶降解RBBR的酶促反应经验模型，该模型对RBBR降解率的预测值与实验实测值具有较好的线性关系，相关系数R2为0.97，能够较为准确的描述和预测漆酶降解RBBR的动力学过程。
漆酶/介体系统可有效降解三苯甲烷类染料酸性品红，反应条件为温度30°C，pH值4.0，酸性品红初始浓度100mg/L，漆酶用量2U/mL，介体HOBT浓度0.05%（w/v）时，反应40min后酸性品红降解率即达到90%以上。根据降解过程中酶促反应速率与时间、温度、染料浓度以及漆酶和介体用量等的关系，建立了白腐菌P. conchatus漆酶降解染料的动力学数学模型，并利用该模型详细研究和考察了漆酶降解染料过程中各反应条件对酶促反应速率的影响。
自制了一种新型气升式生物反应器，该系统可有效进行白腐菌P. conchatus漆酶对多批次染料的连续降解，并通过流动管路利用连续光谱法建立了在线实时监测反应器中染料降解过程的方法。根据多批次RBBR和酸性品红降解过程中染料降解率与时间、反应批次数等的关系，建立了白腐菌P. conchatus漆酶连续降解多批次染料的动力学数学模型，该模型对RBBR和酸性品红降解率的预测值与实验实测值具有较好的线性关系，相关系数R2都能达到0.999以上，能够较为准确的描述和预测漆酶连续降解多批次染料的动力学过程。利用该模型研究和考察了多批次染料连续降解过程中漆酶的重复利用对酶促反应速率的影响，结果表明，漆酶连续降解多批次染料的反应初速度与反应批次数呈线性关系，反应初速度随反应批次的增加有所降低，但是反应初速度下降较慢，表明处理过程中白腐菌P. conchatus漆酶具有较好的稳定性。
Laccase is a multicopper enzyme that can oxidize phenols or aromatic amines by reducingmolecular oxygen to water. This enzyme is remarkably non-specific regarding substrates, andits zone of application can be widened even further in the presence of the aromatic electrontransfer or radical-forming mediators. Laccase has the potential to be implemented in variousindustrial fields, such as pulp biobleaching, discoloration of textile dye, wastewater treatment,polymer syntheses, etc. However, a major limitation for successful industrial application oflaccase in the bioremediation processes mentioned above is the high costs of laccaseproduction. The development of approaches for reducing the costs of laccase production withhigh efficiency and environmental friendliness by optimising the fermentation medium andcondition is a top priority.
In the present paper, the production of laccase by white-rot fungus Panus conchatus insubmerged cultures was investigated. Extracellular laccase formation by P. conchatus can beconsiderably stimulated by the addition of wheat bran and copper sulfate. The maximallaccase activity, approximately196.1U/mL, was obtained in the shake flasks. Submergedculture on larger scale was carried out in a7.5L stirred-tank fermentor, in which the highestactivity of laccase was about200U/mL. The good regression coefficients indicated that theLogistic model was justifiable for the expression and prediction of laccase production by P.conchatus in submerged cultures. Crude laccase of P. conchatus was also used forbiobleaching in LQP1P1sequence with reed pulp. It showed that kappa number, brightness,physical properties and yellowing number of pulp were improved in a certain extent after thebleaching by laccase. The pulp bleachability was increased comparatively after treated bylaccase/mediator system.
Laccase production by P. conchatus in submerged culture using low-cost medium andcooking liquor of bagasse was investigated. Soybean meal and copper sulfate were shown tobe the excellent supporters and played positive roles in enzyme production with the maximallaccase activity of476.7U/mL in the shake flasks. Xylose was found to be the majority of themonosaccharide in liquor from the cooking with diluted H2SO4. The biomass of white-rotfungi Panus conchatus, Flammulina velutipes and Psathyrella candolleana increased during fermentation in the liquid medium based on the cooking liquor without detoxification, butonly F. velutipes showed excellent enzyme production with the maximal laccase activity ofabout200U/L in the medium.
Laccase produced by P. conchatus in submerged culture was purified to electrophoretichomogeneity by precipitation, ultrafiltration, ion-exchange chromatography andsize-exclusion chromatography with a final purification fold of6.77and a yield of74.1%. Theenzyme was found to be a typical fungal laccase with molecular mass of65kDa and Kmvalueof5.7μ2for ABTS. The optimum pH and temperature for laccase activity were2.5and60oC, respectively. The purified laccase retained97.9%of its initial activity after26dinculbation at pH8.0and4oC. The extracellular crude laccase was effective in decolorizationof synthetic dyes, decolorizing95.2%of RBBR in0.5h and84.1%of Methyl Orange in2hwithout redox mediators, approximately100%of Fuchsin Acid in1h with the addition of0.1%HOBT as the meditor.
It was found that anthraquinone dye RBBR could be directely degraded by the crudelaccase of P. conchatus. The degradation of more than90%for100mg/L of RBBR wasobtained in15min with1U/mL of laccase. The relationships between the degradation of dyeand time, pH, temperature, as well as the concentration of laccase were established. Aempirical model of dye degradation was developed based on the experimental results. Thegood regression coefficient indicated that the models were justifiable for the expression andprediction of RBBR degradation.
A detailed investigation concerning to effects of pH, temperature, concentrations of dye,laccase and mediator on the reaction rate of dye degradation by laccase/mediator system(LMS) was conducted. Triphenylmethane dye Fuchsin Acid could be effectively degraded byLMS. The degradation of more than90%for100mg/L Fuchsin Acid was obtained in40minwith2U/mL of laccase and0.5%of HOBT. A kinetic model of reaction rate of dyedegradation by laccase/HOBT system was developed based on the experimental results. Therewas a good agreement between the calculated and measured data, indicating that the modelwas reasonable expression and predication for the reaction rate during dye degradation.
An airlift bioreactor was developed and used to degradation of dye by laccase afterrepeated addition of RBBR or Fuchsin Acid. A Uv-vis spectrophotometric method was established for rapid and temporally resolved monitoring the degradation of dye in thebioreactor. The result showed that the activity of P. conchatus laccase was stable in thebioreactor and had good degradation ability for both of the two dyes. The degradations ofmore than90%and60%of RBBR and Fuchsin Acid were obtained, respectively, in every30min with8times of dye additions. A kinetic model of degradation reaction rate with everytime of dyes addition was developed based on the experimental results. The good regressioncoefficient of the calculated and measured data indicated that the model were justifiable forthe expression and predication for the reaction rate during dye degradation in the bioreactor.
白腐菌Panus conchatus（贝壳状革耳菌）是一种常见的可食用和药用的野生真菌，对木质素的降解具有较强的选择性。本研究将白腐菌P. conchatus漆酶的生产由实验室摇瓶培养扩大到7.5L机械搅拌式发酵罐中，并对搅拌转速、通气量、温度等影响液体深层发酵的关键因素进行了考察和优化。结果表明，麦麸和硫酸铜能够显著提高摇瓶发酵中白腐菌P. conchatus漆酶的产量，缩短发酵周期，发酵液中漆酶活力最高可达196.1U/mL。通气量、培养温度和搅拌转速对白腐菌P. conchatus在机械搅拌式发酵罐中的产酶效率影响较大，最佳发酵条件为通气量1.0vvm，温度30°C，转速300rpm，漆酶活力在发酵22d时达到最大值约200U/mL，保持了和摇瓶发酵相同的产酶水平。Logistic模型能够较为准确的描述和预测白腐菌P. conchatus产漆酶的动力学过程，酶活测定值与方程计算值的线性相关系数R2达到了0.95以上。将该漆酶用于芦苇浆的生物漂白，纸浆经漆酶/介体系统处理后，卡伯值降低，可漂性有所提高，有利于后续漂白工段对纸浆中残余木素的脱除，提高漂白浆的白度。
以农业加工副产物和蔗渣酸法蒸煮废液为基础，研究用于白腐菌培养和漆酶生产的廉价原材料。结果表明，豆粕最适宜作为发酵底物进行漆酶的生产，液体培养基中漆酶最高活力达到476.7U/mL。木糖是蔗渣酸法蒸煮废水中最主要的单糖类物质，白腐菌Panus conchatus，Flammulina velutipes和Psathyrella candolleana都可在未经脱毒处理的废水制备的产酶培养基中生长，其中F. velutipes对废水毒性的抵抗力较强，其生物量和最高漆酶活力分别达到26.3g/L和200U/L。
通过盐析、超滤、离子交换色谱和凝胶过滤色谱等方法，将白腐菌P. conchatus漆酶纯化至电泳纯级别，其比活力达到912.3U/mg，比原始粗酶液中漆酶比活力提高了6.77倍，总酶活得率为74.1%。该漆酶的分子量约为65kDa，纯化后漆酶显蓝色，其紫外可见光谱学特征表明该漆酶为典型的真菌漆酶。漆酶对底物ABTS的米氏常数Km为5.7μ2，最大反应速度Vmax为31.06mM/(QMR)，最适反应温度和pH值分别为60oC和2.5，漆酶活性在4oC，pH8.0的环境中具有较好的稳定性，保存26d后，漆酶活力还能够保持为初始酶活的约97.9%。白腐菌P. conchatus漆酶可直接对蒽醌染料活性亮蓝RBBR和偶氮染料甲基橙进行脱色，粗酶液处理RBBR0.5h后，染料脱色率达到95.2%；处理甲基橙2h后，脱色率为84.1%。在介体HOBT存在的条件下，三苯甲烷类染料酸性品红可被粗酶液在1h内完全脱色。
白腐菌P. conchatus漆酶可有效降解蒽醌染料RBBR，反应条件为温度30°C，pH值4.0，RBBR初始浓度100mg/L，漆酶用量1U/mL时，反应15min后RBBR降解率即达到90%以上。根据降解过程中RBBR降解率与时间、温度、染料浓度以及漆酶用量等的关系，建立了白腐菌P. conchatus漆酶降解RBBR的酶促反应经验模型，该模型对RBBR降解率的预测值与实验实测值具有较好的线性关系，相关系数R2为0.97，能够较为准确的描述和预测漆酶降解RBBR的动力学过程。
漆酶/介体系统可有效降解三苯甲烷类染料酸性品红，反应条件为温度30°C，pH值4.0，酸性品红初始浓度100mg/L，漆酶用量2U/mL，介体HOBT浓度0.05%（w/v）时，反应40min后酸性品红降解率即达到90%以上。根据降解过程中酶促反应速率与时间、温度、染料浓度以及漆酶和介体用量等的关系，建立了白腐菌P. conchatus漆酶降解染料的动力学数学模型，并利用该模型详细研究和考察了漆酶降解染料过程中各反应条件对酶促反应速率的影响。
自制了一种新型气升式生物反应器，该系统可有效进行白腐菌P. conchatus漆酶对多批次染料的连续降解，并通过流动管路利用连续光谱法建立了在线实时监测反应器中染料降解过程的方法。根据多批次RBBR和酸性品红降解过程中染料降解率与时间、反应批次数等的关系，建立了白腐菌P. conchatus漆酶连续降解多批次染料的动力学数学模型，该模型对RBBR和酸性品红降解率的预测值与实验实测值具有较好的线性关系，相关系数R2都能达到0.999以上，能够较为准确的描述和预测漆酶连续降解多批次染料的动力学过程。利用该模型研究和考察了多批次染料连续降解过程中漆酶的重复利用对酶促反应速率的影响，结果表明，漆酶连续降解多批次染料的反应初速度与反应批次数呈线性关系，反应初速度随反应批次的增加有所降低，但是反应初速度下降较慢，表明处理过程中白腐菌P. conchatus漆酶具有较好的稳定性。
Laccase is a multicopper enzyme that can oxidize phenols or aromatic amines by reducingmolecular oxygen to water. This enzyme is remarkably non-specific regarding substrates, andits zone of application can be widened even further in the presence of the aromatic electrontransfer or radical-forming mediators. Laccase has the potential to be implemented in variousindustrial fields, such as pulp biobleaching, discoloration of textile dye, wastewater treatment,polymer syntheses, etc. However, a major limitation for successful industrial application oflaccase in the bioremediation processes mentioned above is the high costs of laccaseproduction. The development of approaches for reducing the costs of laccase production withhigh efficiency and environmental friendliness by optimising the fermentation medium andcondition is a top priority.
In the present paper, the production of laccase by white-rot fungus Panus conchatus insubmerged cultures was investigated. Extracellular laccase formation by P. conchatus can beconsiderably stimulated by the addition of wheat bran and copper sulfate. The maximallaccase activity, approximately196.1U/mL, was obtained in the shake flasks. Submergedculture on larger scale was carried out in a7.5L stirred-tank fermentor, in which the highestactivity of laccase was about200U/mL. The good regression coefficients indicated that theLogistic model was justifiable for the expression and prediction of laccase production by P.conchatus in submerged cultures. Crude laccase of P. conchatus was also used forbiobleaching in LQP1P1sequence with reed pulp. It showed that kappa number, brightness,physical properties and yellowing number of pulp were improved in a certain extent after thebleaching by laccase. The pulp bleachability was increased comparatively after treated bylaccase/mediator system.
Laccase production by P. conchatus in submerged culture using low-cost medium andcooking liquor of bagasse was investigated. Soybean meal and copper sulfate were shown tobe the excellent supporters and played positive roles in enzyme production with the maximallaccase activity of476.7U/mL in the shake flasks. Xylose was found to be the majority of themonosaccharide in liquor from the cooking with diluted H2SO4. The biomass of white-rotfungi Panus conchatus, Flammulina velutipes and Psathyrella candolleana increased during fermentation in the liquid medium based on the cooking liquor without detoxification, butonly F. velutipes showed excellent enzyme production with the maximal laccase activity ofabout200U/L in the medium.
Laccase produced by P. conchatus in submerged culture was purified to electrophoretichomogeneity by precipitation, ultrafiltration, ion-exchange chromatography andsize-exclusion chromatography with a final purification fold of6.77and a yield of74.1%. Theenzyme was found to be a typical fungal laccase with molecular mass of65kDa and Kmvalueof5.7μ2for ABTS. The optimum pH and temperature for laccase activity were2.5and60oC, respectively. The purified laccase retained97.9%of its initial activity after26dinculbation at pH8.0and4oC. The extracellular crude laccase was effective in decolorizationof synthetic dyes, decolorizing95.2%of RBBR in0.5h and84.1%of Methyl Orange in2hwithout redox mediators, approximately100%of Fuchsin Acid in1h with the addition of0.1%HOBT as the meditor.
It was found that anthraquinone dye RBBR could be directely degraded by the crudelaccase of P. conchatus. The degradation of more than90%for100mg/L of RBBR wasobtained in15min with1U/mL of laccase. The relationships between the degradation of dyeand time, pH, temperature, as well as the concentration of laccase were established. Aempirical model of dye degradation was developed based on the experimental results. Thegood regression coefficient indicated that the models were justifiable for the expression andprediction of RBBR degradation.
A detailed investigation concerning to effects of pH, temperature, concentrations of dye,laccase and mediator on the reaction rate of dye degradation by laccase/mediator system(LMS) was conducted. Triphenylmethane dye Fuchsin Acid could be effectively degraded byLMS. The degradation of more than90%for100mg/L Fuchsin Acid was obtained in40minwith2U/mL of laccase and0.5%of HOBT. A kinetic model of reaction rate of dyedegradation by laccase/HOBT system was developed based on the experimental results. Therewas a good agreement between the calculated and measured data, indicating that the modelwas reasonable expression and predication for the reaction rate during dye degradation.
An airlift bioreactor was developed and used to degradation of dye by laccase afterrepeated addition of RBBR or Fuchsin Acid. A Uv-vis spectrophotometric method was established for rapid and temporally resolved monitoring the degradation of dye in thebioreactor. The result showed that the activity of P. conchatus laccase was stable in thebioreactor and had good degradation ability for both of the two dyes. The degradations ofmore than90%and60%of RBBR and Fuchsin Acid were obtained, respectively, in every30min with8times of dye additions. A kinetic model of degradation reaction rate with everytime of dyes addition was developed based on the experimental results. The good regressioncoefficient of the calculated and measured data indicated that the model were justifiable forthe expression and predication for the reaction rate during dye degradation in the bioreactor.
引文
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