纤维素酶降解机理的研究进展
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摘要
纤维素酶在生物炼制过程中具有重要地位,因而十分有必要深度解析酶解作用机理.纤维素是由D-葡萄糖聚合形成的高分子聚合物,它的降解在木质纤维素解聚过程中最为关键,由于其分子结构相当稳定,需要纤维素酶各结构域及组分协同作用,才能将其高效降解为可利用的糖类.本文总结国内外学者对纤维素酶降解机理的最新进展,首先从纤维素酶分子结构与功能的相互作用出发,揭示结构与功能间的互作关系,阐述生物催化分子机制,可为优化酶结构增强酶功能提供理论依据;其次概述纤维素酶降解机理的作用特点,论述酶促协同及其外部环境对纤维素酶活力的重要影响,可为酶解体系优化设计提供借鉴;最后,结合纤维素酶降解机理总结概括了有效降低酶成本的途径.
Cellulase plays an important role in the biorefinery process,so it is necessary to deeply understand the mechanism of enzymatic hydrolysis.Cellulose as polymer compound formed by the polymerization of D-glucose,which is an important component of biomass,and the degradation of cellulose is the most significant step in the destructing process of lignocellulose.Due to its stable molecular structure,only through synergistic action of various domains and components of the cellulase can it be biodegraded into available saccharides.The recent research on the mechanism of cellulase degradation at home and abroad was reviewed.This review firstly explored the interaction between structure and function to reveal the relationship of between them and analyzed the molecular mechanism of biocatalyst,which provides theoretical basis for optimizing cellulase structure to improve catalytic ability.Secondly,the characteristics and molecular structure of cellulase degradation mechanism were discussed,and the important effect of its enzymatic synergy and environment on the enzyme activity was also investigated in order to provide references for the optimization design of enzymatic hydrolysis system.Finally,by combining the mechanism of cellulase biodegradation,the effective ways to reduce enzyme costs was summarized.
Cellulase plays an important role in the biorefinery process,so it is necessary to deeply understand the mechanism of enzymatic hydrolysis.Cellulose as polymer compound formed by the polymerization of D-glucose,which is an important component of biomass,and the degradation of cellulose is the most significant step in the destructing process of lignocellulose.Due to its stable molecular structure,only through synergistic action of various domains and components of the cellulase can it be biodegraded into available saccharides.The recent research on the mechanism of cellulase degradation at home and abroad was reviewed.This review firstly explored the interaction between structure and function to reveal the relationship of between them and analyzed the molecular mechanism of biocatalyst,which provides theoretical basis for optimizing cellulase structure to improve catalytic ability.Secondly,the characteristics and molecular structure of cellulase degradation mechanism were discussed,and the important effect of its enzymatic synergy and environment on the enzyme activity was also investigated in order to provide references for the optimization design of enzymatic hydrolysis system.Finally,by combining the mechanism of cellulase biodegradation,the effective ways to reduce enzyme costs was summarized.
引文
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[3] WILSON D B.Cellulases and biofuels [J].Current Opi-nion in Biotechnology,2009,20(3):295- 299.
[4] RUBIN E M.Genomics of cellulosic biofuels [J].Nature,2008,454(7206):841- 845.
[5] MYERS F L,NORTHCOTE D.Partial purification and some properties of a cellulase from Helix pomatia [J].Biochemical Journal,1959,71(4):749- 756.
[6] REESE E T,SIU R G H,LEVINSON H S.The biological degradation of soluble cellulose derivatives and its relationship to the mechanism of cellulose hydrolysis [J].Journal of Bacteriology,1950,59(4):485.
[7] ZECHEL D L,WITHERS S G.Glycosidase mechanisms:anatomy of a finely tuned catalyst [J].Accounts of Chemi-cal Research,2000,33(1):11- 18.
[8] ARTZI L,BAYER E A,MORAIS S.Cellulosomes:bacterial nanomachines for dismantling plant polysaccharides [J].Nature Reviews Microbiology,2017,15(2):83- 95.
[9] 高培基.纤维素酶降解机制及纤维素酶分子结构与功能研究进展 [J].自然科学进展,2003,13(1):23- 31.GAO Peiji.Adances in cellulase degradation mechanism,structure and function [J].Progress in Natural Science,2003,13(1):23- 31.
[10] VAN TILBEURGH H,TOMME P,CLAEYSSENS M,et al.Limited proteolysis of the cellobiohydrolase I from Trichoderma reesei [J].FEBS letters,1986,204(2):223- 227.
[11] PAYNE C M,KNOTT B C,MAYES H B,et al.Fungal cellulases [J].Chemical reviews,2015,115(3):1308- 1448.
[12] BOMMARIUS A S,SOHN M,KANG Y Z,et al.Protein engineering of cellulases [J].Current Opinion in Biotechnology,2014,29:139- 145.
[13] HALDAR D,SEN D,GAYEN K.A review on the production of fermentable sugars from lignocellulosic biomass through conventional and enzymatic route—a comparison [J].International Journal of Green Energy,2016,13(12):1232- 1253.
[14] QUIROZ-CASTANEDA R E,FOLCH-MALLOL J L.Hydrolysis of biomass mediated by cellulases for the production of sugars[M]//Sustainable degradation of lignocellulosic biomass-techniques,applications and commercialization.London:IntechOpen,2013.
[15] SPEZIO M,WILSON D B,KARPLUS P A.Crystal Structure of the catalytic domain of a Thermophilic endocellulase [J].Biochemistry,1993,32(38):9906- 9916.
[16] 张小梅,李单单,王禄山,等.纤维素酶家族及其催化结构域分子改造的新进展 [J].生物工程学报,2013,29(4):422- 433.ZHANG Xiaomei,LI Dandan,WANG Lushan,et al.Molecular engineering of cellulase catalytic domain based on glycoside hydrolase family [J].Chinese Journal of Bio-technology,2013,29(4):422- 433.
[17] ZHENG F,TU T,WANG X Y,et al.Enhancing the catalytic activity of a novel GH5 cellulase GtCeⅠ5 from Gloeophyllum trabeum CBS 900.73 by site-directed mutagenesis on loop 6 [J].Biotechnology for Biofuels,2018,11(1):6/1- 13.
[18] BORISOVA A S,ENEYSKAYA E V,JANA S,et al.Correlation of structure,function and protein dynamics in GH7 cellobiohydrolases from Trichoderma atroviride,T.reesei and T.harzianum [J].Biotechnology for Biofuels,2018,11(1):5/1- 22.
[19] CHEN M,HIMMEL M E,WILSON D B,et al.Simulation studies of substrate recognition by the exocellulase CelF from Clostridium cellulolyticum [J].Biotechnology and Bioengineering,2016,113(7):1433- 1440.
[20] QIAN M D,GUAN S S,SHAN Y M,et al.Structural and molecular basis of cellulase Cel48F by computational modeling:insight into catalytic and product release mechanism [J].Journal of Structural Biology,2016,194(3):347- 356.
[21] GILKES N,WARREN R,MILLER R,et al.Precise excision of the cellulose binding domains from two Cellulomonas fimi cellulases by a homologous protease and the effect on catalysis [J].Journal of Biological Chemistry,1988,263(21):10401- 10407.
[22] CANTAREL B L,COUTINHO P M,RANCUREL C,et al.The carbohydrate-active enzymes database (CAZy):an expert resource for glycogenomics [J].Nucleic Acids Research,2008,37(suppl_1):D233-D238.
[23] IGARASHI K,KOIVULA A,WADA M,et al.High speed atomic force microscopy visualizes processive movement of Trichoderma reesei cellobiohydrolase I on crystalline cellulose [J].Journal of Biological Chemistry,2009,284(52):36186- 36190.
[24] CARRARD G,KOIVULA A,SODERLUND H,et al.Cellulose-binding domains promote hydrolysis of different sites on crystalline cellulose [J].Proceedings of the National Academy of Sciences,2000,97(19):10342- 10347.
[25] MCCARTNEY L,GILBERT H J,BOLAM D N,et al.Glycoside hydrolase carbohydrate-binding modules as molecular probes for the analysis of plant cell wall polymers [J].Analytical Biochemistry,2004,326(1):49- 54.
[26] DIN N,DAMUDE H G,GILKES N R,et al.C1-Cx revisited:intramolecular synergism in a cellulase [J].Proceedings of the National Academy of Sciences,1994,91(24):11383- 11387.
[27] CATTANEO C,CESARO P,SPERTINO S,et al.Enhanced features of Dictyoglomus turgidum cellulase A engineered with carbohydrate binding module 11 from Clostridium thermocellum [J].Scientific Reports,2018,8(1):4402.
[28] ZHANG Y,YANG F,HU F H,et al.Binding preference of family 1 carbohydrate binding module on nanocrystalline cellulose and nanofibrillar cellulose films assessed by quartz crystal microbalance [J].Cellulose,2018,25(6):3327- 3337.
[29] ZHANG K D,LI W,WANG Y F,et al.Processive degradation of crystalline cellulose by a multimodular endoglucanase via a wirewalking mode [J].Biomacromolecules,2018,19(5):1686- 1696.
[30] LEE J-H,KIM K-N,CHOI Y-J.Identification and charac-terization of a novel inulin binding module (IBM) from the CFTase of Bacillus macerans CFC1 [J].FEMS Microbiology Letters,2004,234(1):105- 110.
[31] HALL M,BANSAL P,LEE J H,et al.Biological pretreatment of cellulose:enhancing enzymatic hydrolysis rate using cellulose-binding domains from cellulases [J].Bioresource Technology,2011,102(3):2910- 2915.
[32] GILKES N,HENRISSAT B,KILBURN D,et al.Domains in microbial β- 1,4-glycanases:sequence conservation,function,and enzyme families [J].Microbiological Reviews,1991,55(2):303- 315.
[33] SRISODSUK M,REINIKAINEN T,PENTTILA M,et al.Role of the interdomain linker peptide of Trichoderma reesei cellobiohydrolase I in its interaction with crystalline cellulose [J].Journal of Biological Chemistry,1993,268(28):20756- 20761.
[34] RECEVEUR V,CZJZEK M,SCHULEIN M,et al.Dimension,shape,and conformational flexibility of a two domain fungal cellulase in solution probed by small angle X-ray scattering [J].Journal of Biological Chemistry,2002,277(43):40887- 40892.
[35] LI P P,ZHOU Y,LI Q,et al.Extending the linker region increases the activity of the Bacillus subtilis cellulase CelI15 [J].Biotechnology Letters,2016,38(9):1587- 1593.
[36] PETKUN S,GRINBERG I R,LAMED R,et al.Reassembly and co-crystallization of a family 9 processive endoglucanase from its component parts:structural and functional significance of the intermodular linker [J].PeerJ,2015,3:e1126.
[37] RUIZ D M,TUROWSKI V R,MURAKAMI M T.Effects of the linker region on the structure and function of modular GH5 cellulases [J].Scientific Reports,2016,6:28504.
[38] FEDAROVICH A,NICHOLAS R A,DAVIES C.Unusual conformation of the SxN motif in the crystal structure of penicillin-binding protein A from Mycobacterium tuberculosis [J].Journal of Molecular Biology,2010,398(1):54- 65.
[39] WU B,ZHENG S,PEDROSO M M,et al.Processivity and enzymatic mechanism of a multifunctional family 5 endoglucanase from Bacillus subtilis BS- 5 with potential applications in the saccharification of cellulosic substrates [J].Biotechnology for Biofuels,2018,11(1):20/1- 15.
[40] KIM I J,LEE H J,CHOI I G,et al.Synergistic proteins for the enhanced enzymatic hydrolysis of cellulose by cellulase [J].Applied Microbiology Biotechnology,2014,98(20):8469- 8480.
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