生物质白腐菌改性与抗性变化的关系研究
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摘要
由于能源危机日益严重，利用木质纤维素类生物质获取生物基产品成为研究热点。通过纤维素酶水解反应从生物质中得到发酵底物——糖类是获取生物基产品的物质基础，但是在天然木质纤维素中，纤维素的低可及度与结晶性、木质素与半纤维素的屏障效应、木质素对纤维素酶的非有效吸附、酶的吸附与解吸附、反应过程的非均相性等诸多因素导致生物质对纤维素酶水解具有很强的抗性，成为生物质有效利用的瓶颈。预处理是减弱抗性，有效释放酶水解底物，提高生物质利用效率的重要途径。诸多白腐菌对木质素具有强降解能力的特性为生物预处理技术发展开辟了新空间。因此，本论文以建立生物质白腐菌改性体系及明确其与抗性变化的内在联系及作用规律为主要内容进行研究。主要研究结果如下：
以玉米秸秆、稻草、竹粉三种生物质为原料，研究了来自神农架自然保护区典型生境的33株担子菌对生物质改性的效果，结果发现不同生物学属性的担子菌对生物质的改性效果差异显著，不同担子菌菌株与生物质类型之间需要组配。适合玉米秸秆、稻草、竹粉三种生物质的菌株依次为菌株CD2、Pleurotus sp.4号、Coriolusversicolor。通过改变担子菌降解环境与条件可优化改性效果，与其它组合及对照相比，菌株CD2改性玉米秸秆效果显著，改性后玉米秸秆疏松、颜色变浅。纤维素酶酶解后糖化率高达62．0％，比未处理的玉米秸秆原料提高231．6％，由此建立了具有应用前景和研究价值的“CD2-玉米秸秆”固体发酵改性体系。对菌株CD2进行分子鉴定，确定其分类地位为：担子菌门、伞菌亚门、伞菌纲、多孔菌目(无褶菌目)、齿耳菌科、耙菌属Irpex lacteus。该菌株在PDA与玉米秸秆基质上最适生长温度不同，分别为30℃与28℃。
在不同降解阶段，I.lacteus CD2通过对木质纤维素不同组分的选择性降解对玉米秸秆进行改性、腐朽。早期选择性降解半纤维素，中期选择性降解与修饰木质素，后期同步降解三种木质纤维素组分。改性与腐朽过程中木质纤维素超分子官能团发生变化的主要类型为醚键水解转化、脱甲基、开环降解等。对腐朽过程中木质素降解酶活性及胞外酶液铁还原能力研究表明：腐朽早、中期未分泌胞外木质素降解酶，但具有铁还原能力，后期胞外滤液具有较低的LiP酶活性。另外，该菌株具有胞壁漆酶。木质素选择性迅速降解时期无胞外木质素降解酶活性，表明I.lacteus CD2降解木质素具有非胞外木质素降解酶主导的特殊机制。
为明确I.lacteus CD2改性玉米秸秆后酶解糖化率显著提高的基质特征，对不同处理后的玉米秸秆样品进行酶解反应过程动力学与酶解负荷动力学研究发现，基于Bailey模型提出的“转化率-酶负荷”公式能够很好地描述转化率随酶负荷的变化规律，拟合参数最大转化率P_1可用于表征生物质底物酶解转化的潜力。以生物质中不能被转化的能源底物的比例定义酶解抗性，依托P_1建立了生物质酶解抗性的定量评价方法。量化的酶解抗性与I.lacteus CD2预处理后玉米秸秆糖化率呈负相关，说明该定量指标能够科学、全面地表征生物质酶解的基质特征。
进一步通过组分测定、XRD、N_2吸附法、ESEM等手段，从I.lacteus CD2改性过程中木质素修饰与含量变化、半纤维素含量变化等化学障碍解除，结晶度、比表面积、孔径分布、表面微观结构等物理屏障变化等多个角度，研究了I.lacteus CD2降低玉米秸秆酶解抗性的机制。结果表明：多种因素共同促使I.lacteus CD2降低酶解抗性，包括木质素修饰、降解，半纤维素含量降低，比表面积、孔隙度增加，菌丝对植物组织的穿透及细胞壁的腐蚀。其中，酸不溶性木质素含量、酸溶性木质素含量分别与酶解抗性具有一定程度负相关与正相关性，表明木质素降解程度对生物质酶解抗性降低起主要作用，结晶度变化对木质纤维素酶解抗性影响不大。该研究不仅揭示了白腐菌生物质改性与酶解抗性降低的内在联系，也为深入进行生物质白腐菌预处理应用研究提供了理论及技术支撑。
Due to the increasingly serious energy crisis and environment problem, the development of bio-based industrial products from lignocellulosic biomass is in great demand, for example, cellulosic ethanol. For production of chemicals from biomass, enzymatic hydrolysis of cellulose and hemicellulose to fermentable sugars is generally employed. However, the conversion efficiency of natural materials is usually very poor, low efficiency of sugar molecule releasing. Biomass recalcitrance, a general designation of substrate-related factors, is responsible for the efficiency of sugar molecule releasing. Many factors contributing to high recalcitrance of biomass mainly include structural heterogeneity and complexity of cell wall, precisely arranged crystalline cellulose, low accessibility of cellulose, lignification and the heterogeneous reaction of saccharification. Of these characteristics above, which are the most important still remains unclear. Pretreatment is one of the key points to enhance the efficiency of biomass utilization and relieve recalcitrance. Due to low energy-consuming and environment friendly, biological pretreatment has received extensive attention. White rot fungi, most belonging to basidiomycetes, are the most promising microorganisms for biological pretreatment due to their capacity of degrading lignin to water and carbon dioxide. Therefore, biomass degradation system by white rot fungi was firstly established, and then the relationship between recalcitrance change and biodegradation was investigated in this paper.
To establish a high efficient degradation system, 33 strains of basidiomycetes, obtained from Shennongjia Nature Reserve in China, were employed to treat three kinds of biomass. After enzymatic hydrolysis of pretreated biomass, transformation rates and saccharification rates were used to evaluate the effect of different pretreatments. The result indicated that the effect of different degradation system varied a lot, with the suitable strains for pretreatment of corn stover, wheat straw and bamboo were strain CD2, Pleurotus sp. No.4 and Coriolus versicolor, respectively. The effect of biological pretreatment could be optimized by altering the cultivation conditions and medium for biodegradation by basidiomycetes. Strain CD2 showed the greatest superiority in promoting enzymatic hydrolysis, with the highest saccharification rate reaching 62.0%, which was 2.32 times of untreated corn stover. Therefore, the best biodegradation system of "strain CD2-corn stover", which was the most promising biopretreatment for cellulosic ethanol production, was chosen for further study. The strain CD2, identified as Irpex lacteus by ITS sequence analysis, is a white rot fungus. The optimum growth temperature was respectively 30℃and 28℃on PDA and corn stover medium. Corn stover pretreated with I. lacteus CD2 was brickie and turned white from its original color of brown.
Corn stover was decayed by different mode with I. lacteus CD2 at varying durations. Hemicellulose and lignin was respectively selectively degraded at the initial and middle stage of biodegradation, with the content of acid soluble lignin being significantly enhanced. Cellulose, hemicellulose and lignin were simultaneously degraded at the last stage. During biodegradation and modification, ether hydrolysis, demethylation and cleavage of aromatic rings were the main changes on lignocellulose function groups. No extracellular ligninolytic enzymes were detected during 120 days of cultivation on corn stover medium except for low LiP activity after 25 days. In PDB medium, low level of mycelium-associated laccase, intracellular MnP and LiP could be detected. FeRA could be detected in the extracellular fermentation extracts all along the decay periods. The results suggested that lignin degradation by I. lacteus CD2 followed a special mechanism independent on extracellular ligninases.
To find the key characteristics influencing saccharification rate, studies on kinetics of enzymatic hydrolysis under different enzyme loading for varying reaction time were carried out. The results indicated initial velocity could not represent the final efficient of enzymatic hydrolysis. Transformation rates of corn stover samples versus enzyme loading followed the same curve proposed by Bailey. Based on this, biomass recalcitrance to enzyme hydrolysis was quantified and calculated as the percentage of holocellulose not hydrolyzed to reducing sugars during enzymatic hydrolysis. The quantified recalcitrance was proved to be in negative correlation with saccharification rate, which illustrated that this parameter could scientifically estimate the general characteristic of biomass for enzymatic hydrolysis.
To make certain the chemical and physical mechanism of the reduced recalcitrance by I. lacteus CD2, further research mainly focused on the changes of lignin, hemicellulose, crystallinity index, surface area, pore size distribution, and surface structure, which were developed by means of components' content determination, XRD analysis, nitrogen adsorption method and ESEM analysis. The results indicated that many factors contributed to the reduced recalcitrance to enzymatic hydrolysis, including lignin degradation and modification, hemicellulose degradation, increased surface area and porosity, mycelium penetration. Acid insoluble lignin and acid soluble lignin were respectively in positive and negative correlation with recalcitrance, which demonstrated that the reduced recalcitrance by I. lacteus CD2 was mainly determined by the extent of lignin degradation and modification. The crystallinity index of cellulose didn't significantly influence recalcitrance. This thesis not only proclaimed the inherent correlation between reduced recalcitrance and biomass quality altering by I. lacteus CD2, but also provided theoretical and technical foundation for biological pretreatment of biomass with white rot fungi.
以玉米秸秆、稻草、竹粉三种生物质为原料，研究了来自神农架自然保护区典型生境的33株担子菌对生物质改性的效果，结果发现不同生物学属性的担子菌对生物质的改性效果差异显著，不同担子菌菌株与生物质类型之间需要组配。适合玉米秸秆、稻草、竹粉三种生物质的菌株依次为菌株CD2、Pleurotus sp.4号、Coriolusversicolor。通过改变担子菌降解环境与条件可优化改性效果，与其它组合及对照相比，菌株CD2改性玉米秸秆效果显著，改性后玉米秸秆疏松、颜色变浅。纤维素酶酶解后糖化率高达62．0％，比未处理的玉米秸秆原料提高231．6％，由此建立了具有应用前景和研究价值的“CD2-玉米秸秆”固体发酵改性体系。对菌株CD2进行分子鉴定，确定其分类地位为：担子菌门、伞菌亚门、伞菌纲、多孔菌目(无褶菌目)、齿耳菌科、耙菌属Irpex lacteus。该菌株在PDA与玉米秸秆基质上最适生长温度不同，分别为30℃与28℃。
在不同降解阶段，I.lacteus CD2通过对木质纤维素不同组分的选择性降解对玉米秸秆进行改性、腐朽。早期选择性降解半纤维素，中期选择性降解与修饰木质素，后期同步降解三种木质纤维素组分。改性与腐朽过程中木质纤维素超分子官能团发生变化的主要类型为醚键水解转化、脱甲基、开环降解等。对腐朽过程中木质素降解酶活性及胞外酶液铁还原能力研究表明：腐朽早、中期未分泌胞外木质素降解酶，但具有铁还原能力，后期胞外滤液具有较低的LiP酶活性。另外，该菌株具有胞壁漆酶。木质素选择性迅速降解时期无胞外木质素降解酶活性，表明I.lacteus CD2降解木质素具有非胞外木质素降解酶主导的特殊机制。
为明确I.lacteus CD2改性玉米秸秆后酶解糖化率显著提高的基质特征，对不同处理后的玉米秸秆样品进行酶解反应过程动力学与酶解负荷动力学研究发现，基于Bailey模型提出的“转化率-酶负荷”公式能够很好地描述转化率随酶负荷的变化规律，拟合参数最大转化率P_1可用于表征生物质底物酶解转化的潜力。以生物质中不能被转化的能源底物的比例定义酶解抗性，依托P_1建立了生物质酶解抗性的定量评价方法。量化的酶解抗性与I.lacteus CD2预处理后玉米秸秆糖化率呈负相关，说明该定量指标能够科学、全面地表征生物质酶解的基质特征。
进一步通过组分测定、XRD、N_2吸附法、ESEM等手段，从I.lacteus CD2改性过程中木质素修饰与含量变化、半纤维素含量变化等化学障碍解除，结晶度、比表面积、孔径分布、表面微观结构等物理屏障变化等多个角度，研究了I.lacteus CD2降低玉米秸秆酶解抗性的机制。结果表明：多种因素共同促使I.lacteus CD2降低酶解抗性，包括木质素修饰、降解，半纤维素含量降低，比表面积、孔隙度增加，菌丝对植物组织的穿透及细胞壁的腐蚀。其中，酸不溶性木质素含量、酸溶性木质素含量分别与酶解抗性具有一定程度负相关与正相关性，表明木质素降解程度对生物质酶解抗性降低起主要作用，结晶度变化对木质纤维素酶解抗性影响不大。该研究不仅揭示了白腐菌生物质改性与酶解抗性降低的内在联系，也为深入进行生物质白腐菌预处理应用研究提供了理论及技术支撑。
Due to the increasingly serious energy crisis and environment problem, the development of bio-based industrial products from lignocellulosic biomass is in great demand, for example, cellulosic ethanol. For production of chemicals from biomass, enzymatic hydrolysis of cellulose and hemicellulose to fermentable sugars is generally employed. However, the conversion efficiency of natural materials is usually very poor, low efficiency of sugar molecule releasing. Biomass recalcitrance, a general designation of substrate-related factors, is responsible for the efficiency of sugar molecule releasing. Many factors contributing to high recalcitrance of biomass mainly include structural heterogeneity and complexity of cell wall, precisely arranged crystalline cellulose, low accessibility of cellulose, lignification and the heterogeneous reaction of saccharification. Of these characteristics above, which are the most important still remains unclear. Pretreatment is one of the key points to enhance the efficiency of biomass utilization and relieve recalcitrance. Due to low energy-consuming and environment friendly, biological pretreatment has received extensive attention. White rot fungi, most belonging to basidiomycetes, are the most promising microorganisms for biological pretreatment due to their capacity of degrading lignin to water and carbon dioxide. Therefore, biomass degradation system by white rot fungi was firstly established, and then the relationship between recalcitrance change and biodegradation was investigated in this paper.
To establish a high efficient degradation system, 33 strains of basidiomycetes, obtained from Shennongjia Nature Reserve in China, were employed to treat three kinds of biomass. After enzymatic hydrolysis of pretreated biomass, transformation rates and saccharification rates were used to evaluate the effect of different pretreatments. The result indicated that the effect of different degradation system varied a lot, with the suitable strains for pretreatment of corn stover, wheat straw and bamboo were strain CD2, Pleurotus sp. No.4 and Coriolus versicolor, respectively. The effect of biological pretreatment could be optimized by altering the cultivation conditions and medium for biodegradation by basidiomycetes. Strain CD2 showed the greatest superiority in promoting enzymatic hydrolysis, with the highest saccharification rate reaching 62.0%, which was 2.32 times of untreated corn stover. Therefore, the best biodegradation system of "strain CD2-corn stover", which was the most promising biopretreatment for cellulosic ethanol production, was chosen for further study. The strain CD2, identified as Irpex lacteus by ITS sequence analysis, is a white rot fungus. The optimum growth temperature was respectively 30℃and 28℃on PDA and corn stover medium. Corn stover pretreated with I. lacteus CD2 was brickie and turned white from its original color of brown.
Corn stover was decayed by different mode with I. lacteus CD2 at varying durations. Hemicellulose and lignin was respectively selectively degraded at the initial and middle stage of biodegradation, with the content of acid soluble lignin being significantly enhanced. Cellulose, hemicellulose and lignin were simultaneously degraded at the last stage. During biodegradation and modification, ether hydrolysis, demethylation and cleavage of aromatic rings were the main changes on lignocellulose function groups. No extracellular ligninolytic enzymes were detected during 120 days of cultivation on corn stover medium except for low LiP activity after 25 days. In PDB medium, low level of mycelium-associated laccase, intracellular MnP and LiP could be detected. FeRA could be detected in the extracellular fermentation extracts all along the decay periods. The results suggested that lignin degradation by I. lacteus CD2 followed a special mechanism independent on extracellular ligninases.
To find the key characteristics influencing saccharification rate, studies on kinetics of enzymatic hydrolysis under different enzyme loading for varying reaction time were carried out. The results indicated initial velocity could not represent the final efficient of enzymatic hydrolysis. Transformation rates of corn stover samples versus enzyme loading followed the same curve proposed by Bailey. Based on this, biomass recalcitrance to enzyme hydrolysis was quantified and calculated as the percentage of holocellulose not hydrolyzed to reducing sugars during enzymatic hydrolysis. The quantified recalcitrance was proved to be in negative correlation with saccharification rate, which illustrated that this parameter could scientifically estimate the general characteristic of biomass for enzymatic hydrolysis.
To make certain the chemical and physical mechanism of the reduced recalcitrance by I. lacteus CD2, further research mainly focused on the changes of lignin, hemicellulose, crystallinity index, surface area, pore size distribution, and surface structure, which were developed by means of components' content determination, XRD analysis, nitrogen adsorption method and ESEM analysis. The results indicated that many factors contributed to the reduced recalcitrance to enzymatic hydrolysis, including lignin degradation and modification, hemicellulose degradation, increased surface area and porosity, mycelium penetration. Acid insoluble lignin and acid soluble lignin were respectively in positive and negative correlation with recalcitrance, which demonstrated that the reduced recalcitrance by I. lacteus CD2 was mainly determined by the extent of lignin degradation and modification. The crystallinity index of cellulose didn't significantly influence recalcitrance. This thesis not only proclaimed the inherent correlation between reduced recalcitrance and biomass quality altering by I. lacteus CD2, but also provided theoretical and technical foundation for biological pretreatment of biomass with white rot fungi.
引文
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