细菌Caldicellulosiruptor bescii嗜热纤维素酶系功能及协同作用研究
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摘要
当前，石油资源日益枯竭，能源消耗日趋增多。寻求可再生能源以降低对不可再生资源依赖以及应对气候变化，已经成为国际社会关注的焦点。木质纤维素生物质作为世界上最多的可再生有机物，被认为是生物燃料和其他增值产物最主要的来源。为将木质纤维素生物质转变为重要的工业产品，必须将其转化成可发酵的单糖或寡糖，纤维素酶在此过程中具有关键作用。嗜热纤维素酶与常温纤维素酶相比具有诸多优势，如热处理木质纤维素时酶稳定性好，因而对反应体系具有良好的兼容性。开发来源于嗜热菌的新型纤维素酶及探讨其功能及对不溶纤维素的协同水解机理是重要的研究课题。本论文克隆、异源表达了来源于真细菌Caldicellulosiruptor bescii的嗜热渐进性内切纤维素酶、纤维二糖水解酶和β葡萄糖苷酶，并对其底物特异性、产物特异性及其协同作用水解纤维素等进行了系统地研究。
来源于嗜热真细菌C. bescii的CbCelA是模块化的酶，由糖苷水解酶第9家族结构域（GH9）和第48家族结构域（GH48）通过3个糖类结合蛋白（CBM3c和两个重复的CBM3b）连接。为研究每个模块的功能特点，我们克隆构建了一系列CbCelA截短蛋白（CbCel9A_3cbm、CbCel9A_2cbm、CbCel9A_1cbm、CbCel9A、CbCbh48A、2cbm_CbCbh48A、CBM3c和CBM3b）。通过测定各重组蛋白底物特异性、产物特异性、吸附能力和协同能力等特征，揭示各模块水解纤维素的功能。结果表明CbCel9A为渐进性内切纤维素酶，对不溶性纤维素具有较高活力；CbCbh48A为纤维二糖水解酶，对固体纤维素活力很低，但是它可以通过协同作用加速CbCel9A_1cbm或其他内切纤维素酶水解活力。紧邻的CBM3c对CbCel9A活力、热稳定性及渐进性具有重要作用，且帮助CbCel9A吸附不溶底物。CBM3b则帮助CbCel9A_1cbm和CbCbh48A吸附到固体底物上，提高水解不溶底物活力和渐进性。
细菌β葡萄糖苷酶作为纤维素酶系统的一个主要成分，负责水解纤维二糖等纤维寡糖为葡萄糖。为了获得新型耐热β葡萄糖苷酶有效用于水解纤维素，我们克隆、异源表达了来源于嗜热真细菌C. bescii的β葡萄糖苷酶CbBgl1A（GenBank：ACM59590.1）。通过系统地测定其底物特异性、催化动力学、结构建模和分子对接，以及与其他嗜热纤维素酶协同作用。我们发现CbBgl1A可以在高温下水解一系列纤维寡糖，β类二糖，和芳基-β-糖苷；高浓度底物不能够抑制其活力，对产物葡萄糖有较高的耐受性，且可以与外切纤维素酶和渐进性内切纤维素酶以较高协同性水解固体纤维素。
为进一步深入研究纤维素酶协同水解纤维素，我们选定上述表征的CbCel9A_3cbm和CbCbh48A以及本实验室已有的FnCel5A三种嗜热纤维素酶，通过不同比例组合，在高温下水解再生不定型纤维素RAC，并在反应体系中加入定量的CbBgl1A以消除产物抑制。结果表明CbCel9A_3cbm在水解RAC过程中发挥最重要作用，CbCbh48A作用最小。转化率在最初两个小时最高，随着孵育时间延长，水解速度显著降低。60oC水解24小时后，CbCel9A_3cbm，CbCbh48A，和FnCel5A的最佳比例为60:20:20，转化率达到81.9%。进一步提高反应温度（75oC）后，在2小时内最高转化率可达到46%。
In view of rising prices of crude oil due to increasing fuel demands, the need foralternative sources of bioenergy inresponse to decreasing of non-renewable resourcesand climate change is expected to increase sharply in the coming years.Lignocellulosic biomass, as the most abundant renewable source of organic matter onthe earth, has been identified as the prime source of biofuels and other value-addedproducts. To initiate the production of industrially important products from cellulosicbiomass, bioconversion of the cellulosic components into fermentable sugars isnecessary, whereas cellulase is the key element in this process. Thermostable enzymeshave been studied for their abilities to hydrolyze lignocellulosic materials since theyhave several potential advantages compared to their mesophilic counterparts, such asbetter hydrolytic efficiency and stability at higher temperatures. Screening novelthermostable cellulase and detailed study into the biochemical and catalytic propertiesof them, together with an improved understanding of the synergistic mechanisms inthe presence of other cellulase components is important to assess their full potentialfrom both fundamental and applied standpoints. Hearein we report cloning andoverexpression of processive endoglucanase, cellobiohydrylase and β-glucosidasefrom thermophile Caldicellulosiruptor bescii, we systematically determined thesubstrate specificities, product specificities and synergistic effects of the enzymes.
A bifunctional enzyme designated CbCelA from C. bescii is a modular protein, ithas a glycoside hydrolase family9(GH9) module at the N-terminus and a GH48module at the C-terminus. Located between the two modules are threecarbohydrate-binding modules (CBMs), one of which belongs to CBM3c subfamily,other two modules are repeated one belongs to CBM3b subfamily. To dissect function of the each module, several truncated forms (CbCel9A_3cbm，CbCel9A_2cbm,CbCel9A_1cbm, CbCel9A, CbCbh48A,2cbm_CbCbh48A, CBM3c and CBM3b)were constructed. The biochemical properties were described by determiningsubstrate and product specificities, ability of adsorption on cellulose and synergisticeffects with other cellulases. The results suggested that the GH9domain (CbCel9A)was a processive endoglucanase with high activity on insoluble cellulose, whereasGH48(CbCbh48A) was a cellobiohydrolase. It possessed very low activity towardscellulose but could synergistically work with CbCel9A_1cbm or other cellulase tohydrolyze substrate. The CBM3c module was very important to the activity andthermostability of CbCel9A, meanwhile, this protein helped CbCel9A partially bind tocellulose. The CBM3b module could adsorb to cellulose, resulting in increasedconcentration of enzymes on surface of cellulose, thereby enhancing the activities ofCbCel9A_1cbm and CbCbh48A.
Bacterial β-glucosidase (BGL) is a major component of the cellulase system andis responsible for the hydrolysis of cellobiose and short chain oligosaccharides intoglucose. In an effort to obtain new thermostable BGLs desirable for the hydrolysis ofcellulose, herein we report a novel recombinant BGL from the thermophile C. bescii(CbBgl1A). Enzyme properties of CbBgl1A were determined by analyses of substratespecificity, catalytic kinetics, structure modeling, along with synergistic effects withother cellulase components. We found that CbBgl1A could hydrolyze a range ofcellooligosaccharides, β-diglycosides, as well as aryl-β-glycosides at high temperature.It was resistance to both high concentrations of substrate and product, and its catalysiswas highly synergistic with exocellulases (cellobiohydrolases) and processiveendocellulase in the hydrolysis of cellulose.
To further understand the synergistic mechanisms of cellulases, CbCel9A_3cbmand CbCbh48A, together with another thermalstable endoglucanase FnCel5A whichwas previously characterized in our lab, were selected to form a minimal set ofbacterial cellulases for bioconversion of cellulose. In addition, fixed amounts ofCbBgl1A were added to eliminate product inhibition.21reactions containing differentratios of three cellulases and fixed amounts of CbBgl1A were studied on disordered high-accessibility regenerated amorphous cellulose (RAC). Processive endoglucanaseCbCel9A_3cbm was the most important for high cellulose digestibility, whereasCbCbh48A contributed the least. The optimal ratio for maximum cellulosedigestibility (81.9%) was60:20:20(CbCel9A_3cbm: CbCbh48A: FnCel5A) after24h incubation at60oC. The digestibility improved along with temperature increase.After2h incubation at75oC, the best mixture for maximum digestibility (46%) was80:0:20, which was2.82-fold improvement than that of CbCel9A alone.
来源于嗜热真细菌C. bescii的CbCelA是模块化的酶，由糖苷水解酶第9家族结构域（GH9）和第48家族结构域（GH48）通过3个糖类结合蛋白（CBM3c和两个重复的CBM3b）连接。为研究每个模块的功能特点，我们克隆构建了一系列CbCelA截短蛋白（CbCel9A_3cbm、CbCel9A_2cbm、CbCel9A_1cbm、CbCel9A、CbCbh48A、2cbm_CbCbh48A、CBM3c和CBM3b）。通过测定各重组蛋白底物特异性、产物特异性、吸附能力和协同能力等特征，揭示各模块水解纤维素的功能。结果表明CbCel9A为渐进性内切纤维素酶，对不溶性纤维素具有较高活力；CbCbh48A为纤维二糖水解酶，对固体纤维素活力很低，但是它可以通过协同作用加速CbCel9A_1cbm或其他内切纤维素酶水解活力。紧邻的CBM3c对CbCel9A活力、热稳定性及渐进性具有重要作用，且帮助CbCel9A吸附不溶底物。CBM3b则帮助CbCel9A_1cbm和CbCbh48A吸附到固体底物上，提高水解不溶底物活力和渐进性。
细菌β葡萄糖苷酶作为纤维素酶系统的一个主要成分，负责水解纤维二糖等纤维寡糖为葡萄糖。为了获得新型耐热β葡萄糖苷酶有效用于水解纤维素，我们克隆、异源表达了来源于嗜热真细菌C. bescii的β葡萄糖苷酶CbBgl1A（GenBank：ACM59590.1）。通过系统地测定其底物特异性、催化动力学、结构建模和分子对接，以及与其他嗜热纤维素酶协同作用。我们发现CbBgl1A可以在高温下水解一系列纤维寡糖，β类二糖，和芳基-β-糖苷；高浓度底物不能够抑制其活力，对产物葡萄糖有较高的耐受性，且可以与外切纤维素酶和渐进性内切纤维素酶以较高协同性水解固体纤维素。
为进一步深入研究纤维素酶协同水解纤维素，我们选定上述表征的CbCel9A_3cbm和CbCbh48A以及本实验室已有的FnCel5A三种嗜热纤维素酶，通过不同比例组合，在高温下水解再生不定型纤维素RAC，并在反应体系中加入定量的CbBgl1A以消除产物抑制。结果表明CbCel9A_3cbm在水解RAC过程中发挥最重要作用，CbCbh48A作用最小。转化率在最初两个小时最高，随着孵育时间延长，水解速度显著降低。60oC水解24小时后，CbCel9A_3cbm，CbCbh48A，和FnCel5A的最佳比例为60:20:20，转化率达到81.9%。进一步提高反应温度（75oC）后，在2小时内最高转化率可达到46%。
In view of rising prices of crude oil due to increasing fuel demands, the need foralternative sources of bioenergy inresponse to decreasing of non-renewable resourcesand climate change is expected to increase sharply in the coming years.Lignocellulosic biomass, as the most abundant renewable source of organic matter onthe earth, has been identified as the prime source of biofuels and other value-addedproducts. To initiate the production of industrially important products from cellulosicbiomass, bioconversion of the cellulosic components into fermentable sugars isnecessary, whereas cellulase is the key element in this process. Thermostable enzymeshave been studied for their abilities to hydrolyze lignocellulosic materials since theyhave several potential advantages compared to their mesophilic counterparts, such asbetter hydrolytic efficiency and stability at higher temperatures. Screening novelthermostable cellulase and detailed study into the biochemical and catalytic propertiesof them, together with an improved understanding of the synergistic mechanisms inthe presence of other cellulase components is important to assess their full potentialfrom both fundamental and applied standpoints. Hearein we report cloning andoverexpression of processive endoglucanase, cellobiohydrylase and β-glucosidasefrom thermophile Caldicellulosiruptor bescii, we systematically determined thesubstrate specificities, product specificities and synergistic effects of the enzymes.
A bifunctional enzyme designated CbCelA from C. bescii is a modular protein, ithas a glycoside hydrolase family9(GH9) module at the N-terminus and a GH48module at the C-terminus. Located between the two modules are threecarbohydrate-binding modules (CBMs), one of which belongs to CBM3c subfamily,other two modules are repeated one belongs to CBM3b subfamily. To dissect function of the each module, several truncated forms (CbCel9A_3cbm，CbCel9A_2cbm,CbCel9A_1cbm, CbCel9A, CbCbh48A,2cbm_CbCbh48A, CBM3c and CBM3b)were constructed. The biochemical properties were described by determiningsubstrate and product specificities, ability of adsorption on cellulose and synergisticeffects with other cellulases. The results suggested that the GH9domain (CbCel9A)was a processive endoglucanase with high activity on insoluble cellulose, whereasGH48(CbCbh48A) was a cellobiohydrolase. It possessed very low activity towardscellulose but could synergistically work with CbCel9A_1cbm or other cellulase tohydrolyze substrate. The CBM3c module was very important to the activity andthermostability of CbCel9A, meanwhile, this protein helped CbCel9A partially bind tocellulose. The CBM3b module could adsorb to cellulose, resulting in increasedconcentration of enzymes on surface of cellulose, thereby enhancing the activities ofCbCel9A_1cbm and CbCbh48A.
Bacterial β-glucosidase (BGL) is a major component of the cellulase system andis responsible for the hydrolysis of cellobiose and short chain oligosaccharides intoglucose. In an effort to obtain new thermostable BGLs desirable for the hydrolysis ofcellulose, herein we report a novel recombinant BGL from the thermophile C. bescii(CbBgl1A). Enzyme properties of CbBgl1A were determined by analyses of substratespecificity, catalytic kinetics, structure modeling, along with synergistic effects withother cellulase components. We found that CbBgl1A could hydrolyze a range ofcellooligosaccharides, β-diglycosides, as well as aryl-β-glycosides at high temperature.It was resistance to both high concentrations of substrate and product, and its catalysiswas highly synergistic with exocellulases (cellobiohydrolases) and processiveendocellulase in the hydrolysis of cellulose.
To further understand the synergistic mechanisms of cellulases, CbCel9A_3cbmand CbCbh48A, together with another thermalstable endoglucanase FnCel5A whichwas previously characterized in our lab, were selected to form a minimal set ofbacterial cellulases for bioconversion of cellulose. In addition, fixed amounts ofCbBgl1A were added to eliminate product inhibition.21reactions containing differentratios of three cellulases and fixed amounts of CbBgl1A were studied on disordered high-accessibility regenerated amorphous cellulose (RAC). Processive endoglucanaseCbCel9A_3cbm was the most important for high cellulose digestibility, whereasCbCbh48A contributed the least. The optimal ratio for maximum cellulosedigestibility (81.9%) was60:20:20(CbCel9A_3cbm: CbCbh48A: FnCel5A) after24h incubation at60oC. The digestibility improved along with temperature increase.After2h incubation at75oC, the best mixture for maximum digestibility (46%) was80:0:20, which was2.82-fold improvement than that of CbCel9A alone.
引文
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