木质纤维素对里氏木霉产纤维素酶的诱导
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摘要
在化石能源日趋短缺与能源需求的日益增长的矛盾形势下，以及化石能源带来的环境问题日益严重背景下，利用木质纤维原料制备燃料乙醇替代传统化石能源的技术成为当今世界的研究热点之一。纤维素酶能够有效将农作物秸秆等富含纤维素的物质水解为单糖，用于乙醇发酵，因此如何获得廉价高效的纤维素酶对我国经济和社会可持续发展具有重要的意义。本论文以里氏木霉RUT C30为产酶菌株，研究了不同碳源诱导产纤维素酶过程。
以农作物秸秆为原料，研究了稀酸处理、碱处理和绿液预处理工艺条件；不同预处理条件对秸秆碳源诱导纤维素酶的影响；廉价产酶碳源和预处理方式确立；通过响应面分析优化确立稀酸处理纸浆产酶培养基；通过预处理秸秆和稀酸处理纸浆碳源单独分批补料以及混合碳源分批补料制备纤维素酶，以延长产酶周期提高产酶效率；使用电镜扫描和X-衍射等手段分析不同原料通过预处理条件提高产酶能力的原因。主要结果如下：
比较了稀酸处理、碱处理和绿液预处理方式对不同秸秆碳源诱导产酶能力，结果表明绿液预处理方式处理条件温和、处理方式简单、原料损失小，诱导产酶能力最佳。比较了绿液预处理对稻草、玉米秸秆、棉秆和麦秆诱导里氏木霉产酶的影响，实验表明麦秆比较适合绿液预处理方式用于产酶碳源。比较了不同绿液预处理条件下木质纤维原料的浆料分析、纤维素酶活力与纤维素酶解得率，结果表明，预处理条件越剧烈，原料损失越大，聚糖回收率越低，而木质素移除率越高。绿液预处理对麦秆的影响程度分别为总碱量、蒸煮温度、硫化度。
麦秆产酶最佳绿液处理条件是蒸煮温度140℃，总碱量4%和硫化度20%，12g/L麦秆纤维素为碳源，间歇培养合成纤维素酶，滤纸酶活第6天最高达4.23FPIU/mL。碳源浓度提高到20g/L麦秆纤维素，滤纸酶活最高达5.89FPIU/mL。
绿液预处理麦秆酶水解的最适宜条件是蒸煮温度150℃，总碱量8%和硫化度40%的条件。当采用10%底物浓度，15FPIU/g纤维素，酶解72h，以商品纤维素酶为列，葡聚糖得率和木聚糖得率分别由10.50%和5.43%提高到61.41%和55.65%。自产纤维素酶酶解效果好于商品纤维素酶，葡聚糖得率和木聚糖得率分别由61.41%和55.65%提高到81.07%和80.02%，提高了32.01%和43.79%。
稀酸处理纸浆为碳源产酶，通过PB实验得到影响纤维素酶合成的三个主要因素为酸处理纸浆浓度、C/N和硫酸镁浓度，经响应面实验优化这三个因素得到最佳结果，当酸处理纸浆浓度30.94g/L，C/N为4.06，硫酸镁浓度0.36g/L，滤纸酶活在120h达到最大为10.35FPIU/mL，此时β-G、蛋白质、酶产率和酶得率分别为4.98IU/mL、3.5g/L、86.25IU/(L·h)和334.52IU/g稀酸纸浆。
通过分批补料产酶方式可以大幅度提高产纤维素酶量。实验得到了以绿液处理麦秆为单独碳源分批补料产酶，最佳补料方式是：补料速度为5.5g/(L·d)纤维素时，第5天开始补料，第10天FPA达到最高值9.56FPIU/mL，此时pH、β-G和酶产率分别为6.56、2.44IU/mL和39.83IU/(L·h)。以稀酸处理纸浆为单独碳源分批补料产酶，最佳补料方式是：补料速度为4g/(L·d)纤维素，第1天开始补料，第10天FPA达到最大值为17.58IU/mL，此时pH、β-G、酶产率和酶得率分别为6.57、7.01IU/mL、73.25IU/(L·h)和399.55IU/g纤维素。以绿液处理麦秆和稀酸处理纸浆混合碳源分批补料产酶，最佳补料方式是，以8.0g/L绿液处理麦秆纤维素为起始碳源，稀酸处理纸浆为补料碳源，补料速度为4g/(L·d)纤维素时，第9天FPA达到最大为15.33FPIU/mL，此时酶产率的酶得率分别为70.97IU/(L·h)和638.75IU/g纤维素，高于间歇产酶和单独碳源分批补料的产酶效率。
Under the circumstance of the contradiction between the shortage of fossil fuel and theincreasing demand for energy, as well as environmental problems become increasingly seriousbring by the fossil energy, using lignocellulose to prepare fuel ethanol instead of traditional fossilenergy has become one the hottest topics for research in today’s world. Cellulase can effectivelyhydrolyze cellulose-rich straws into monosaccharide, which can be fermented to produce fuelethanol. It is very meaningful in the sustaining development of our society and economy that howto produce the costless and effective cellulase. The process of cellulase induction fromTrichoderma reesei RUT C30using different carbon sources was studied.
In this dissertation, straws were used as the raw materials pretreated by dilute acid, alkaliand green liquor. Effects of pretreatment conditions for different carbon source on cellulaseinduction were compared. Cheap carbon source and pretreatment method for enzyme productionwas optimized. Response Surface Analysis was applied to optimize the enzyme productionmedium when dilute-acid treated pulp was used as the carbon sourse. To extend enzymeproduction cycle and to improve the efficiency of enzyme production, pretreated wheat straw anddilute-acid pretreated pulp were used separately or together in batch fermentation or fed-batchfermentation. SEM and X-ray diffraction were used to examine the structure changes oflignocellulose pretreated with different conditions and to investigate the effect on the increase ofthe enzyme production ability. The main results are as follows.
The capability of enzyme induction by different straws pretreated with dilute acid, alkali,and green liquor was compared. Results indicated that green liquor pretreatment was preferredbecause of mild conditions, simple processing, less material loss, and high enzyme inducingability. The effects of green liquor pretreatment of rice straw, corn stalk, cotton stalk and wheatstraw on cellulase production with Trichoderma reesei were studied. Data showed that wheatstraw pretreated by green liquor was the more suitable carbon source for enzyme production.Analysis of pulp yield, cellulase activity and yield of enzymatic hydrolysis with different greenliquid pretreated lignocellulose indicated that higher weight loss, lower glycan recovery, andhigher level of delignification were observed when more severe condition of pretreatment was applied. The order of influence of green liquor pretreatment on wheat straw was total alkalicharge (Na2O, on oven dry material, TTA), cooking temperature, and sulfidity.
For enzyme production with wheat straw, the best green liquor pretreatment condition wascooking temperature140°C, total alkali charge4%, and sulfidity20%. The pretreated wheatstraw was used as carbon source for enzyme production. When12g/L of cellulose was used forcellulase synthesis, batch fermentation could reach a filter paper activity of4.23FPIU/mL at thesixth day. Increasing the carbon source concentration to20g/L of cellulose resulted a filterpaper activity of5.89FPIU/mL.
For enzymatic hydrolysis of wheat straw, the optimum condition of green liquorpretreatment was cooking temperature150℃, total alkali charge8%, and sulfidity40%.Hydrolysis was conducted at substrate concentration of10%for72h with a cellulase loading of15FPIU/g cellulose. Compared to un-pretreated wheat straw, the green liquor pretreated whatstraw was hydrolyzed more efficiently. When commercial cellulose was used, hydrolysis yield ofglucan was increase from10.50%to61.41%, and hydrolysis yield of xylan was increase from5.43%to55.65%. When lab-produced cellulase was used, however, hydrolysis was furtherimproved. Hydrolysis yield of glucan and hydrolysis yield of xylan were increased to81.07%and80.02%, respectively.
Pulp pretreated with1.5%H2SO4was the most efficient carbon source in inducingTrichoderma reesei for cellulase production. Plackett-Burman (PB) experiments indicated thatthe three main factors of pretreatment affecting enzyme synthesis were pulp concentration, C/Nand MgSO4concentration. Response Surface Analysis experiments optimized these three factors.When the dilute acid-hydrolyzed pulp（AHP）concentration was30.94g/L, C/N was4.06, andMgSO4concentration was0.36g/L, after120h growth, the filter paper activity reached it’smaximum of10.35FPIU/mL, with β-glucosidase activity, protein content, cellulase productionrate and cellulase yield being4.98IU/mL,3.5g/L,86.25IU/L·h and334.52IU/g cellulose,respectively.
Fed-batch culture could increase cellulase production significantly and was optimized fordifferent carbon sources. When green liquor pretreated wheat straw was used as the carbonsource in fed-batch enzyme production, the optimal feeding condition was addition of5.5g/L·dcellulose starting at the fifth day. At the tenth day filter paper activity reached the highest valueof9.56FPIU/mL; pH, β-glucosidase activity and cellulase production rate were6.56,2.44IU/mL and39.83IU/L·h, respectively. In fed-batch enzyme production with dilute-acid pretreatedpulp as the single carbon source, the best feeding mode was adding4g/L·d cellulose every dayfrom the first day. At the tenth day filter paper activity reached the highest value of17.58FPIU/mL, whereas pH, β-glucosidase activity, cellulase production rate and cellulose yield were6.57,7.01IU/mL,73.25IU/L·h and399.55IU/g cellulose, respectively. When green liquidpretreated wheat straw and dilute-acid pretreated pulp were used as mixed carbon source for fed-batch enzyme production, the best way of carbon source feeding was started with greenliquid pretreated wheat straw at8.0g/L of cellulose, followed by addition of dilute-acidpretreated pulp, from the first day and on, at a concentration of4g/L·d cellulose. The highestvalue of filter paper activity,15.33FPIU/mL, was determined at the ninth day. At the meantimecellulase production rate and cellulose yield were70.97g/L·h and638.75IU/g cellulose,respectively, higher than that when single carbon source was used.
以农作物秸秆为原料，研究了稀酸处理、碱处理和绿液预处理工艺条件；不同预处理条件对秸秆碳源诱导纤维素酶的影响；廉价产酶碳源和预处理方式确立；通过响应面分析优化确立稀酸处理纸浆产酶培养基；通过预处理秸秆和稀酸处理纸浆碳源单独分批补料以及混合碳源分批补料制备纤维素酶，以延长产酶周期提高产酶效率；使用电镜扫描和X-衍射等手段分析不同原料通过预处理条件提高产酶能力的原因。主要结果如下：
比较了稀酸处理、碱处理和绿液预处理方式对不同秸秆碳源诱导产酶能力，结果表明绿液预处理方式处理条件温和、处理方式简单、原料损失小，诱导产酶能力最佳。比较了绿液预处理对稻草、玉米秸秆、棉秆和麦秆诱导里氏木霉产酶的影响，实验表明麦秆比较适合绿液预处理方式用于产酶碳源。比较了不同绿液预处理条件下木质纤维原料的浆料分析、纤维素酶活力与纤维素酶解得率，结果表明，预处理条件越剧烈，原料损失越大，聚糖回收率越低，而木质素移除率越高。绿液预处理对麦秆的影响程度分别为总碱量、蒸煮温度、硫化度。
麦秆产酶最佳绿液处理条件是蒸煮温度140℃，总碱量4%和硫化度20%，12g/L麦秆纤维素为碳源，间歇培养合成纤维素酶，滤纸酶活第6天最高达4.23FPIU/mL。碳源浓度提高到20g/L麦秆纤维素，滤纸酶活最高达5.89FPIU/mL。
绿液预处理麦秆酶水解的最适宜条件是蒸煮温度150℃，总碱量8%和硫化度40%的条件。当采用10%底物浓度，15FPIU/g纤维素，酶解72h，以商品纤维素酶为列，葡聚糖得率和木聚糖得率分别由10.50%和5.43%提高到61.41%和55.65%。自产纤维素酶酶解效果好于商品纤维素酶，葡聚糖得率和木聚糖得率分别由61.41%和55.65%提高到81.07%和80.02%，提高了32.01%和43.79%。
稀酸处理纸浆为碳源产酶，通过PB实验得到影响纤维素酶合成的三个主要因素为酸处理纸浆浓度、C/N和硫酸镁浓度，经响应面实验优化这三个因素得到最佳结果，当酸处理纸浆浓度30.94g/L，C/N为4.06，硫酸镁浓度0.36g/L，滤纸酶活在120h达到最大为10.35FPIU/mL，此时β-G、蛋白质、酶产率和酶得率分别为4.98IU/mL、3.5g/L、86.25IU/(L·h)和334.52IU/g稀酸纸浆。
通过分批补料产酶方式可以大幅度提高产纤维素酶量。实验得到了以绿液处理麦秆为单独碳源分批补料产酶，最佳补料方式是：补料速度为5.5g/(L·d)纤维素时，第5天开始补料，第10天FPA达到最高值9.56FPIU/mL，此时pH、β-G和酶产率分别为6.56、2.44IU/mL和39.83IU/(L·h)。以稀酸处理纸浆为单独碳源分批补料产酶，最佳补料方式是：补料速度为4g/(L·d)纤维素，第1天开始补料，第10天FPA达到最大值为17.58IU/mL，此时pH、β-G、酶产率和酶得率分别为6.57、7.01IU/mL、73.25IU/(L·h)和399.55IU/g纤维素。以绿液处理麦秆和稀酸处理纸浆混合碳源分批补料产酶，最佳补料方式是，以8.0g/L绿液处理麦秆纤维素为起始碳源，稀酸处理纸浆为补料碳源，补料速度为4g/(L·d)纤维素时，第9天FPA达到最大为15.33FPIU/mL，此时酶产率的酶得率分别为70.97IU/(L·h)和638.75IU/g纤维素，高于间歇产酶和单独碳源分批补料的产酶效率。
Under the circumstance of the contradiction between the shortage of fossil fuel and theincreasing demand for energy, as well as environmental problems become increasingly seriousbring by the fossil energy, using lignocellulose to prepare fuel ethanol instead of traditional fossilenergy has become one the hottest topics for research in today’s world. Cellulase can effectivelyhydrolyze cellulose-rich straws into monosaccharide, which can be fermented to produce fuelethanol. It is very meaningful in the sustaining development of our society and economy that howto produce the costless and effective cellulase. The process of cellulase induction fromTrichoderma reesei RUT C30using different carbon sources was studied.
In this dissertation, straws were used as the raw materials pretreated by dilute acid, alkaliand green liquor. Effects of pretreatment conditions for different carbon source on cellulaseinduction were compared. Cheap carbon source and pretreatment method for enzyme productionwas optimized. Response Surface Analysis was applied to optimize the enzyme productionmedium when dilute-acid treated pulp was used as the carbon sourse. To extend enzymeproduction cycle and to improve the efficiency of enzyme production, pretreated wheat straw anddilute-acid pretreated pulp were used separately or together in batch fermentation or fed-batchfermentation. SEM and X-ray diffraction were used to examine the structure changes oflignocellulose pretreated with different conditions and to investigate the effect on the increase ofthe enzyme production ability. The main results are as follows.
The capability of enzyme induction by different straws pretreated with dilute acid, alkali,and green liquor was compared. Results indicated that green liquor pretreatment was preferredbecause of mild conditions, simple processing, less material loss, and high enzyme inducingability. The effects of green liquor pretreatment of rice straw, corn stalk, cotton stalk and wheatstraw on cellulase production with Trichoderma reesei were studied. Data showed that wheatstraw pretreated by green liquor was the more suitable carbon source for enzyme production.Analysis of pulp yield, cellulase activity and yield of enzymatic hydrolysis with different greenliquid pretreated lignocellulose indicated that higher weight loss, lower glycan recovery, andhigher level of delignification were observed when more severe condition of pretreatment was applied. The order of influence of green liquor pretreatment on wheat straw was total alkalicharge (Na2O, on oven dry material, TTA), cooking temperature, and sulfidity.
For enzyme production with wheat straw, the best green liquor pretreatment condition wascooking temperature140°C, total alkali charge4%, and sulfidity20%. The pretreated wheatstraw was used as carbon source for enzyme production. When12g/L of cellulose was used forcellulase synthesis, batch fermentation could reach a filter paper activity of4.23FPIU/mL at thesixth day. Increasing the carbon source concentration to20g/L of cellulose resulted a filterpaper activity of5.89FPIU/mL.
For enzymatic hydrolysis of wheat straw, the optimum condition of green liquorpretreatment was cooking temperature150℃, total alkali charge8%, and sulfidity40%.Hydrolysis was conducted at substrate concentration of10%for72h with a cellulase loading of15FPIU/g cellulose. Compared to un-pretreated wheat straw, the green liquor pretreated whatstraw was hydrolyzed more efficiently. When commercial cellulose was used, hydrolysis yield ofglucan was increase from10.50%to61.41%, and hydrolysis yield of xylan was increase from5.43%to55.65%. When lab-produced cellulase was used, however, hydrolysis was furtherimproved. Hydrolysis yield of glucan and hydrolysis yield of xylan were increased to81.07%and80.02%, respectively.
Pulp pretreated with1.5%H2SO4was the most efficient carbon source in inducingTrichoderma reesei for cellulase production. Plackett-Burman (PB) experiments indicated thatthe three main factors of pretreatment affecting enzyme synthesis were pulp concentration, C/Nand MgSO4concentration. Response Surface Analysis experiments optimized these three factors.When the dilute acid-hydrolyzed pulp（AHP）concentration was30.94g/L, C/N was4.06, andMgSO4concentration was0.36g/L, after120h growth, the filter paper activity reached it’smaximum of10.35FPIU/mL, with β-glucosidase activity, protein content, cellulase productionrate and cellulase yield being4.98IU/mL,3.5g/L,86.25IU/L·h and334.52IU/g cellulose,respectively.
Fed-batch culture could increase cellulase production significantly and was optimized fordifferent carbon sources. When green liquor pretreated wheat straw was used as the carbonsource in fed-batch enzyme production, the optimal feeding condition was addition of5.5g/L·dcellulose starting at the fifth day. At the tenth day filter paper activity reached the highest valueof9.56FPIU/mL; pH, β-glucosidase activity and cellulase production rate were6.56,2.44IU/mL and39.83IU/L·h, respectively. In fed-batch enzyme production with dilute-acid pretreatedpulp as the single carbon source, the best feeding mode was adding4g/L·d cellulose every dayfrom the first day. At the tenth day filter paper activity reached the highest value of17.58FPIU/mL, whereas pH, β-glucosidase activity, cellulase production rate and cellulose yield were6.57,7.01IU/mL,73.25IU/L·h and399.55IU/g cellulose, respectively. When green liquidpretreated wheat straw and dilute-acid pretreated pulp were used as mixed carbon source for fed-batch enzyme production, the best way of carbon source feeding was started with greenliquid pretreated wheat straw at8.0g/L of cellulose, followed by addition of dilute-acidpretreated pulp, from the first day and on, at a concentration of4g/L·d cellulose. The highestvalue of filter paper activity,15.33FPIU/mL, was determined at the ninth day. At the meantimecellulase production rate and cellulose yield were70.97g/L·h and638.75IU/g cellulose,respectively, higher than that when single carbon source was used.
引文
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