植物细胞壁抗降解屏障研究进展与展望

doi:10.11707/j.1001-7488.LYKX20220329

摘要/Abstract

摘要：

纤维素是地球上最丰富的可再生资源，经酶促水解反应生成葡萄糖是纤维素转化的理想方式，但转化效率低，探究相关的抑制因素及其特点成为纤维素高值化应用的首要基础科学问题。本研究首先从宏观形态、显微结构、超微结构、分子、基团、元素、化学键和基因水平，多角度、多层次总结制约纤维素酶促水解反应效率的关键抑制因素，阐述抗降解屏障的内涵；其次明确抗降解屏障表现出的在不同生物质的特有异质性、在植物发育不同阶段的高度动态性以及在预处理过程的复杂联动性；最后展望生物质抗降解屏障破解研究的新趋势和新策略：细胞壁界面抗降解屏障的新认识，细胞壁修饰与改造方法，细胞壁精准解构与组分分级利用的新技术和多酶协同酶解体系的建立等。

关键词: 抗降解屏障, 木质纤维素生物质, 生物转化, 酶促水解反应

Abstract:

Cellulose is deemed as the most abundant and renewable resource on the earth. Enzymatic saccharification of cellulose to glucose is an ideal bioconversion, but its low efficiency significantly impedes the development of cellulose industrialization. Recognition and deconstruction of different recalcitrant factors and improving enzymatic hydrolysis efficiency are still the vital scientific problems in the high valorization of cellulose. In this article, various recalcitrant factors that impeding the enzymatic hydrolysis were summarized at multidimensional levels, in the aspects of macrostructure, microstructure, ultra-structure, molecule, chemical group, chemical element, chemical bonds and gene. Furthermore, we revealed that recalcitrant factors exhibited typical heterogeneity in specific biomass, high dynamics in different development stage of plant, and complex relation during the pretreatment. Finally, several novel research directions and strategy on recalcitrant barrier were proposed: cell wall interface recalcitrance, modification and valorization of cell wall from a biorefinery concept and saccharification hydrolysis with synergies of multiple enzymes.

Key words: recalcitrance, lignocellulosic biomass, bioconversion&shy, enzymatic hydrolysis

中图分类号:

鲁彦,李嘉祺,马雨萱,薛慧婷,李冠华. 植物细胞壁抗降解屏障研究进展与展望[J]. 林业科学, 2024, 60(3): 160-168.

Yan Lu,Jiaqi Li,Yuxuan Ma,Huiting Xue,Guanhua Li. Recent Progress on Recalcitrance of Biomass[J]. Scientia Silvae Sinicae, 2024, 60(3): 160-168.

图/表 3

图1

表1

表2

植物细胞壁的基因修饰"

基因/酶 Gene/enzyme	生物质 Plant species	生物学作用 Biological function	参考文献 Reference
GAUT4	柳枝稷Panicum virgatum 美洲黑杨 Populus deltoides 稻Oryza sativa	合成果胶，促进木质素-碳水化合物交联，降低细胞壁糖的可萃取性 Synthesizing pectin, increasing wall calcium and boron, promoting lignin-carbohydrate cross-linkages and decreasing extractability of cell wall sugars	Biswal et al., 2018; Li et al., 2019
OsMYB103L-RNAi	稻Oryza sativa	导致纤维素微纤维破碎无序，降低CrI和DP Leading to a broken and disordered cellulose microfibers and reducing CrI and DP values	Wu et al., 2021
PtoDET2	毛白杨Populus tomentosa	参与木质素合成;提高纤维素CrI和DP；增加半纤维素Xyl/Ara比例 Participating in lignin synthesis; reducing cellulose CrI, DP, hemicellulose Xyl/Ara ratio	Fan et al., 2020a
OsCAD2	稻Oryza sativa	合成木质素；减少H木质素，增加G木质 Synthesizing lignin; reducing H monomer and increasing G monomer levels	Zhang et al., 2021
PtoLAC14	拟南芥 Arabidopsis thaliana	合成木质素；降低S/G比例 Synthesizing lignin; reducing the proportion of S/G	Qin et al., 2020
CAD	稻Oryza sativa	参与木质素的合成和运输 Participating in lignin synthesis and transportation	Martin et al., 2019
OSSUS3	稻Oryza sativa	促进纤维素和半纤维素沉积 Enhancing cellulose and hemicelluloses deposition	Fan et al., 2020b
AtCESA6	稻Oryza sativa	通过促进细胞生长和次级细胞壁沉积；增加纤维素产量 Increasing cellulose production by enhancing cell growth and secondary cell wall deposition	Ai et al., 2021
COBL2	拟南芥Arabidopsis thaliana	增加结晶纤维素的沉积；提高阿拉伯糖和半乳糖的含量 Resulting in the deposition of crystalline cellulose; increasing contents of arabinose and galactose	Ben-Tov et al., 2018
QUA2	拟南芥Arabidopsis thaliana	促进果胶的生物合成和管壁积累，增加纤维素在管壁的定向沉积 Promoting the biosynthesis of pectic HG and accumulation in the wall , increasing the aligned deposition of cellulose in the wall	Du et al., 2020
AtCESA1 ^T166A	拟南芥Arabidopsis thaliana	增加果胶甲基酯化; 降低纤维素合成/沉积 Causing the increase of pectin methylesterification and defects in cellulose synthesis/deposition	Zhang et al., 2022
OsCESA9	稻Oryza sativa	降低纤维素CrI和DP，降低半纤维素Xyl/Ara比例，提高果胶中糖醛酸的含量 Reducing cellulose CrI and DP values and hemicellulose Xyl/Ara ratio, increasing uronic acids from pectin	Yu et al., 2022
PgGUX	白云杉Picea glauca	引入GlcA修饰半纤维Introducing GlcA to decorate hemicellulose	Lyczakowski et al., 2017

表2

参考文献 0

	胡惠仁, 石淑兰, 冯文英. 芦苇与荻木素结构及对制浆性能的影响. 纤维素科学与技术, 2000, 8 (2): 23- 29. doi: 10.3969/j.issn.1004-8405.2000.02.005
	Hu H R, Shi S L, Feng W Y. Comparison of the lignin structures and pulpabilities between reed and amur silver grass. Journal of Cellulose Science and Technology, 2000, 8 (2): 23- 29. doi: 10.3969/j.issn.1004-8405.2000.02.005
	彭霄鹏, 聂双喜, 刘　璟, 等. 基于微波液化的木质纤维素组分分离和转化——纤维素组分的理化和酶解性质. 林业科学, 2019, 55 (5): 134- 141. doi: 10.11707/j.1001-7488.20190515
	Peng X P, Nie S X, Liu J, et al. Physicochemical characterization and enzymatic hydrolysis of cellulosic component fractionated from microwave liquefied lignocellulosic biomass. Scientia Silvae Sinicae, 2019, 55 (5): 134- 141. doi: 10.11707/j.1001-7488.20190515
	李　丹, 苑　琳, 李　猛, 等. 甜菜渣纤维素乙醇研究进展与展望. 生物工程学报, 2016, 32 (7): 880- 888.
	Li D, Yuan L, Li M, et al. Progress on cellulosic ethanol produced from beet pulp. Chinese Journal of Biotechnology, 2016, 32 (7): 880- 888.
	石　宁, 唐文勇, 唐石云, 等. 木质纤维素衍生平台化学品制备液态烷烃的研究进展. 化工进展, 2019, 38 (7): 3097- 3110.
	Shi N, Tang W Y, Tang S Y, et al. Advances in the catalytic conversion of lignocellulosic derived platform chemicals into liquid alkanes. Chemical Industry and Engineering Progress, 2019, 38 (7): 3097- 3110.
	Adani F, Papa G, Schievano A, et al. Nanoscale structure of the cell wall protecting cellulose from enzyme attack. Environmental Science & Technology, 2011, 45 (3): 1107- 1113.
	Aguilera-Segura S M, Di Renzo F, Mineva T. Molecular insight into the cosolvent effect on lignin-cellulose adhesion. Langmuir:the ACS Journal of Surfaces and Colloids, 2020, 36 (47): 14403- 14416. doi: 10.1021/acs.langmuir.0c02794
	Ai Y H, Feng S Q, Wang Y M, et al. Integrated genetic and chemical modification with rice straw for maximum bioethanol production. Industrial Crops and Products, 2021, 173, 114133. doi: 10.1016/j.indcrop.2021.114133
	Amos R A, Mohnen D. Critical review of plant cell wall matrix polysaccharide glycosyltransferase activities verified by heterologous protein expression. Frontiers in Plant Science, 2019, 10, 915. doi: 10.3389/fpls.2019.00915
	Arantes V, Saddler J N. Cellulose accessibility limits the effectiveness of minimum cellulase loading on the efficient hydrolysis of pretreated lignocellulosic substrates. Biotechnology for Biofuels, 2011, 4, 3. doi: 10.1186/1754-6834-4-3
	Avci U, Zhou X L, Pattathil S, et al. Effects of extractive ammonia pretreatment on the ultrastructure and glycan composition of corn stover. Frontiers in Energy Research, 2019, 7, 85. doi: 10.3389/fenrg.2019.00085
	Ben-Tov D, Idan-Molakandov A, Hugger A, et al. The role of COBRA-LIKE 2 function, as part of the complex network of interacting pathways regulating Arabidopsis seed mucilage polysaccharide matrix organization. The Plant Journal:for Cell and Molecular Biology, 2018, 94 (3): 497- 512. doi: 10.1111/tpj.13871
	Biswal A K, Atmodjo M A, Li M, et al. Sugar release and growth of biofuel crops are improved by downregulation of pectin biosynthesis. Nature Biotechnology, 2018, 36, 249- 257. doi: 10.1038/nbt.4067
	Bruno-Soares A M, Matos T J S, Cadima J. Nutritive value of Cistus salvifolius shrubs for small ruminants. Animal Feed Science and Technology, 2011, 165 (3/4): 167- 175.
	Cajnko M M, Oblak J, Grilc M, et al. Enzymatic bioconversion process of lignin: mechanisms, reactions and kinetics. Bioresource Technology, 2021, 340, 125655. doi: 10.1016/j.biortech.2021.125655
	Carpita N C, McCann M C. Redesigning plant cell walls for the biomass-based bioeconomy. Journal of Biological Chemistry, 2020, 295 (44): 15144- 15157. doi: 10.1074/jbc.REV120.014561
	Chen Z, Reznicek W D, Wan C X. Deep eutectic solvent pretreatment enabling full utilization of switchgrass. Bioresource Technology, 2018, 263, 40- 48. doi: 10.1016/j.biortech.2018.04.058
	Collucci D, Bueno R C A, Milagres A M F, et al. Sucrose content, lignocellulose accumulation and in vitro digestibility of sugarcane internodes depicted in relation to internode maturation stage and Saccharum genotypes. Industrial Crops and Products, 2019, 139, 111543. doi: 10.1016/j.indcrop.2019.111543
	Crowe J D, Feringa N, Pattathil S, et al. Identification of developmental stage and anatomical fraction contributions to cell wall recalcitrance in switchgrass. Biotechnology for Biofuels, 2017, 10, 184. doi: 10.1186/s13068-017-0870-5
	DeMartini J D, Pattathil S, Miller J S, et al. Investigating plant cell wall components that affect biomass recalcitrance in poplar and switchgrass. Energy & Environmental Science, 2013, 6 (3): 898.
	dos Santos A C, Ximenes E, Kim Y, et al. Lignin-enzyme interactions in the hydrolysis of lignocellulosic biomass. Trends in Biotechnology, 2019, 37 (5): 518- 531. doi: 10.1016/j.tibtech.2018.10.010
	Du J, Kirui A, Huang S X, et al. Mutations in the pectin methyltransferase QUASIMODO2 influence cellulose biosynthesis and wall integrity in Arabidopsis. The Plant Cell, 2020, 32 (11): 3576- 3597. doi: 10.1105/tpc.20.00252
	Fan C F, Wang G Y, Wu L M, et al. Distinct cellulose and callose accumulation for enhanced bioethanol production and biotic stress resistance in OsSUS3 transgenic rice. Carbohydrate Polymers, 2020a, 232, 115448. doi: 10.1016/j.carbpol.2019.115448
	Fan C F, Yu H, Qin S F, et al. Brassinosteroid overproduction improves lignocellulose quantity and quality to maximize bioethanol yield under green-like biomass process in transgenic poplar. Biotechnology for Biofuels, 2020b, 13, 9. doi: 10.1186/s13068-020-1652-z
	Gomide F T F, da Silva A S, da Silva Bon E P, et al. Modification of microcrystalline cellulose structural properties by ball-milling and ionic liquid treatments and their correlation to enzymatic hydrolysis rate and yield. Cellulose, 2019, 26 (12): 7323- 7335. doi: 10.1007/s10570-019-02578-8
	Gouker F E, Fabio E S, Serapiglia M J, et al. Yield and biomass quality of shrub willow hybrids in differing rotation lengths and spacing designs. Biomass and Bioenergy, 2021, 146, 105977. doi: 10.1016/j.biombioe.2021.105977
	Gu H Q, An R X, Bao J. Pretreatment refining leads to constant particle size distribution of lignocellulose biomass in enzymatic hydrolysis. Chemical Engineering Journal, 2018, 352, 198- 205. doi: 10.1016/j.cej.2018.06.145
	Guo X H, Fu Y J, Miao F W, et al. Efficient separation of functional xylooligosaccharide, cellulose and lignin from poplar via thermal acetic acid/sodium acetate hydrolysis and subsequent kraft pulping. Industrial Crops and Products, 2020, 153, 112575. doi: 10.1016/j.indcrop.2020.112575
	Gupta A, Das S P, Ghosh A, et al. Bioethanol production from hemicellulose rich Populus nigra involving recombinant hemicellulases from Clostridium thermocellum. Bioresource Technology, 2014, 165, 205- 213. doi: 10.1016/j.biortech.2014.03.132
	He J, Huang C X, Lai C H, et al. Elucidation of structure-inhibition relationship of monosaccharides derived pseudo-lignin in enzymatic hydrolysis. Industrial Crops and Products, 2018, 113, 368- 375. doi: 10.1016/j.indcrop.2018.01.046
	He J, Huang C X, Lai C H, et al. Revealing the mechanism of lignin re-polymerization inhibitor in acidic pretreatment and its impact on enzymatic hydrolysis. Industrial Crops and Products, 2022, 179, 114631. doi: 10.1016/j.indcrop.2022.114631
	Himmel M E, Ding S Y, Johnson D K, et al. Biomass recalcitrance: engineering plants and enzymes for biofuels production. Science, 2007, 315 (5813): 804- 807. doi: 10.1126/science.1137016
	Holopainen-Mantila U, Marjamaa K, Merali Z, et al. Impact of hydrothermal pre-treatment to chemical composition, enzymatic digestibility and spatial distribution of cell wall polymers. Bioresource Technology, 2013, 138, 156- 162. doi: 10.1016/j.biortech.2013.03.152
	Hu S W, Wu L M, Persson S, et al. Sweet sorghum and Miscanthus: two potential dedicated bioenergy crops in China. Journal of Integrative Agriculture, 2017, 16 (6): 1236- 1243. doi: 10.1016/S2095-3119(15)61181-9
	Hu Z, Wang Y M, Liu J Y, et al. Integrated NIRS and QTL assays reveal minor mannose and galactose as contrast lignocellulose factors for biomass enzymatic saccharification in rice. Biotechnology for Biofuels, 2021, 14 (1): 1- 13. doi: 10.1186/s13068-020-01854-1
	Huang J F, Xia T, Li G H, et al. Overproduction of native endo-β-1, 4-glucanases leads to largely enhanced biomass saccharification and bioethanol production by specific modification of cellulose features in transgenic rice. Biotechnology for Biofuels, 2019, 12, 11. doi: 10.1186/s13068-018-1351-1
	Isikgor F H, Becer C R. Lignocellulosic biomass: a sustainable platform for the production of bio-based chemicals and polymers. Polymer Chemistry, 2015, 6 (25): 4497- 4559. doi: 10.1039/C5PY00263J
	Jaafar Z, Mazeau K, Boissiére A, et al. Meaning of xylan acetylation on xylan-cellulose interactions: a quartz crystal microbalance with dissipation (QCM-D) and molecular dynamic study. Carbohydrate Polymers, 2019, 226, 115315. doi: 10.1016/j.carbpol.2019.115315
	Karthikeyan O P, Visvanathan C. Bio-energy recovery from high-solid organic substrates by dry anaerobic bio-conversion processes: a review. Reviews in Environmental Science and Bio/Technology, 2013, 12 (3): 257- 284. doi: 10.1007/s11157-012-9304-9
	Kozlova L, Petrova A, Chernyad’ev A, et al. On the origin of bast fiber dislocations in flax. Industrial Crops and Products, 2022, 176, 114382. doi: 10.1016/j.indcrop.2021.114382
	Lee S Y, Sankaran R, Chew K W, et al. Waste to bioenergy: a review on the recent conversion technologies. BMC Energy, 2019, 1 (1): 1- 22. doi: 10.1186/s42500-019-0003-8
	Li G H, He W, Yuan L. Aqueous ammonia pretreatment of sugar beet pulp for enhanced enzymatic hydrolysis. Bioprocess and Biosystems Engineering, 2017, 40 (11): 1603- 1609. doi: 10.1007/s00449-017-1816-9
	Li G H, Sun Y X, Guo W J, et al. Comparison of various pretreatment strategies and their effect on chemistry and structure of sugar beet pulp. Journal of Cleaner Production, 2018, 181, 217- 223. doi: 10.1016/j.jclepro.2018.01.259
	Li G H, Zhang Y, Zhao C, et al. Chemical variation in cell wall of sugar beet pulp caused by aqueous ammonia pretreatment influence enzymatic digestibility of cellulose. Industrial Crops and Products, 2020, 155 (7): 112786.
	Li M, Yoo C G, Pu Y Q, et al. Downregulation of pectin biosynthesis gene GAUT4 leads to reduced ferulate and lignin-carbohydrate cross-linking in switchgrass. Communications Biology, 2019, 2, 22. doi: 10.1038/s42003-018-0265-6
	Liang W L, Liao J S, Qi J R, et al. Physicochemical characteristics and functional properties of high methoxyl pectin with different degree of esterification. Food Chemistry, 2022, 375, 131806. doi: 10.1016/j.foodchem.2021.131806
	Lin C Y, Donohoe B S, Bomble Y J, et al. Iron incorporation both intra- and extra-cellularly improves the yield and saccharification of switchgrass (Panicum virgatum L. ) biomass. Biotechnology for Biofuels, 2021, 14 (1): 55. doi: 10.1186/s13068-021-01891-4
	Lin Y Y, Chen W H, Colin B, et al. Thermodegradation characterization of hardwoods and softwoods in torrefaction and transition zone between torrefaction and pyrolysis. Fuel, 2022, 310, 122281. doi: 10.1016/j.fuel.2021.122281
	Ling Z, Tang W, Su Y, et al. Stepwise allomorphic transformations by alkaline and ethylenediamine treatments on bamboo crystalline cellulose for enhanced enzymatic digestibility. Industrial Crops and Products, 2022, 177, 114450. doi: 10.1016/j.indcrop.2021.114450
	Liu M R, Wang L, Si M Y, et al. New insight into enzymatic hydrolysis of the rice straw and poplar: an in-depth statistical analysis on the multiscale recalcitrance. BioEnergy Research, 2019a, 12 (1): 21- 33. doi: 10.1007/s12155-019-9959-y
	Liu Y R, Nie Y, Lu X M, et al. Cascade utilization of lignocellulosic biomass to high-value products. Green Chemistry, 2019b, 21 (13): 3499- 3535. doi: 10.1039/C9GC00473D
	Liu Y, Xue H T, Miao C Y, et al. Microstructural and histochemical variations of Korshinsk peashrub during aqueous ammonia pretreatment and its impacts on enzymatic digestibility of cellulose. Journal of Environmental Chemical Engineering, 2022, 10 (6): 108613. doi: 10.1016/j.jece.2022.108613
	Lyczakowski J J, Wicher K B, Terrett O M, et al. Removal of glucuronic acid from xylan is a strategy to improve the conversion of plant biomass to sugars for bioenergy. Biotechnology for Biofuels, 2017, 10, 224. doi: 10.1186/s13068-017-0902-1
	Machado G, Leon S, Santos F, et al. Literature review on furfural production from lignocellulosic biomass. Natural Resources, 2016, 7 (3): 115- 129. doi: 10.4236/nr.2016.73012
	Mankar A R, Pandey A, Modak A, et al. Pretreatment of lignocellulosic biomass: a review on recent advances. Bioresource Technology, 2021, 334, 125235. doi: 10.1016/j.biortech.2021.125235
	Mansfield S D, Mooney C, Saddler J N. Substrate and enzyme characteristics that limit cellulose hydrolysis. Biotechnology Progress, 1999, 15 (5): 804- 816. doi: 10.1021/bp9900864
	Martin A F, Tobimatsu Y, Kusumi R, et al. Altered lignocellulose chemical structure and molecular assembly in cinnamyl alcohol dehydrogenase-deficient rice. Scientific Reports, 2019, 9, 17153. doi: 10.1038/s41598-019-53156-8
	Martínez-Abad A, Jiménez-Quero A, Wohlert J, et al. Influence of the molecular motifs of mannan and xylan populations on their recalcitrance and organization in spruce softwoods. Green Chemistry, 2020, 22 (12): 3956- 3970. doi: 10.1039/D0GC01207F
	Mason P J, Furtado A, Marquardt A, et al. Variation in sugarcane biomass composition and enzymatic saccharification of leaves, internodes and roots. Biotechnology for Biofuels, 2020, 13 (1): 201. doi: 10.1186/s13068-020-01837-2
	Negrão D R, Grandis A, Buckeridge M S, et al. Inorganics in sugarcane bagasse and straw and their impacts for bioenergy and biorefining: a review. Renewable and Sustainable Energy Reviews, 2021, 148, 111268. doi: 10.1016/j.rser.2021.111268
	Pielhop T, Reinhard C, Hecht C, et al. Application potential of a carbocation scavenger in autohydrolysis and dilute acid pretreatment to overcome high softwood recalcitrance. Biomass and Bioenergy, 2017, 105, 164- 173. doi: 10.1016/j.biombioe.2017.07.005
	Proietti S, Moscatello S, Fagnano M, et al. Chemical composition and yield of rhizome biomass of Arundo donax L. grown for biorefinery in the Mediterranean environment. Biomass and Bioenergy, 2017, 107, 191- 197. doi: 10.1016/j.biombioe.2017.10.003
	Qin S F, Fan C F, Li X H, et al. LACCASE14 is required for the deposition of guaiacyl lignin and affects cell wall digestibility in poplar. Biotechnology for Biofuels, 2020, 13 (1): 197. doi: 10.1186/s13068-020-01843-4
	Rigual V, Santos T M, Domínguez J C, et al. Evaluation of hardwood and softwood fractionation using autohydrolysis and ionic liquid microwave pretreatment. Biomass and Bioenergy, 2018, 117, 190- 197. doi: 10.1016/j.biombioe.2018.07.014
	Seki Y, Kikuchi Y, Yoshimoto R, et al. Promotion of crystalline cellulose degradation by expansins from Oryza sativa. Planta, 2015, 241 (1): 83- 93. doi: 10.1007/s00425-014-2163-6
	Shen Z H, Zhang K J, Si M Y, et al. Synergy of lignocelluloses pretreatment by sodium carbonate and bacterium to enhance enzymatic hydrolysis of rice straw. Bioresource Technology, 2018, 249, 154- 160. doi: 10.1016/j.biortech.2017.10.008
	Shi J, Pattathil S, Parthasarathi R, et al. Impact of engineered lignin composition on biomass recalcitrance and ionic liquid pretreatment efficiency. Green Chemistry, 2016, 18, 4884- 4895. doi: 10.1039/C6GC01193D
	Sun Y C, Liu X N, Wang T T, et al. Green process for extraction of lignin by the microwave-assisted ionic liquid approach: toward biomass biorefinery and lignin characterization. ACS Sustainable Chemistry & Engineering, 2019, 7 (15): 13062- 13072.
	Suota M J, da Silva T A, Zawadzki S F, et al. Chemical and structural characterization of hardwood and softwood LignoForce™ lignins. Industrial Crops and Products, 2021, 173, 114138. doi: 10.1016/j.indcrop.2021.114138
	Terrett O M, Lyczakowski J J, Yu L, et al. Molecular architecture of softwood revealed by solid-state NMR. Nature Communications, 2019, 10, 4978. doi: 10.1038/s41467-019-12979-9
	Thite V S, Nerurkar A S. Crude xylanases and pectinases from Bacillus spp. along with commercial cellulase formulate an efficient tailor-made cocktail for sugarcane bagasse saccharification. BioEnergy Research, 2020, 13 (1): 286- 300. doi: 10.1007/s12155-019-10050-5
	Tiwari Y M, Sarangi S K. Characterization of raw and alkali treated cellulosic Grewia Flavescens natural fiber. International Journal of Biological Macromolecules, 2022, 209, 1933- 1942. doi: 10.1016/j.ijbiomac.2022.04.169
	Turumtay H. Cell wall engineering by heterologous expression of cell wall-degrading enzymes for better conversion of lignocellulosic biomass into biofuels. BioEnergy Research, 2015, 8 (4): 1574- 1588. doi: 10.1007/s12155-015-9624-z
	Valenzuela-Riffo F, Parra-Palma C, Ramos P, et al. Molecular and structural insights into FaEXPA5, an alpha-expansin protein related with cell wall disassembly during ripening of strawberry fruit. Plant Physiology and Biochemistry, 2020, 154, 581- 589. doi: 10.1016/j.plaphy.2020.06.010
	Valenzuela-Riffo F, Ramos P, Morales-Quintana L. Computational study of FaEXPA1, a strawberry alpha expansin protein, through molecular modeling and molecular dynamics simulation studies. Computational Biology and Chemistry, 2018, 76, 79- 86. doi: 10.1016/j.compbiolchem.2018.05.018
	Wang Y F, Xu X Y, Xue H T, et al. Physical-chemical properties of cell wall interface significantly correlated to the complex recalcitrance of corn straw. Biotechnology for Biofuels, 2021, 14 (1): 196. doi: 10.1186/s13068-021-02047-0
	Wei J H, Wang L Z, Zhai S C, et al. Depolymerization behaviors of naked oat stem cell wall during autohydrolysis in subcritical water. Industrial Crops and Products, 2021, 170, 113679. doi: 10.1016/j.indcrop.2021.113679
	Wu L M, Zhang M L, Zhang R, et al. Down-regulation of OsMYB103L distinctively alters beta-1, 4-glucan polymerization and cellulose microfibers assembly for enhanced biomass enzymatic saccharification in rice. Biotechnology for Biofuels, 2021, 14 (1): 245. doi: 10.1186/s13068-021-02093-8
	Yang B, Voiniciuc C, Fu L B, et al. TRM4 is essential for cellulose deposition in Arabidopsis seed mucilage by maintaining cortical microtubule organization and interacting with CESA3. The New Phytologist, 2019, 221 (2): 881- 895. doi: 10.1111/nph.15442
	Yang Z, Foston M B, O’Neill H, et al. Structural reorganization of noncellulosic polymers observed in situ during dilute acid pretreatment by small-angle neutron scattering. ACS Sustainable Chemistry & Engineering, 2022, 10 (1): 314- 322.
	Yu H, Hu M, Hu Z, et al. Insights into pectin dominated enhancements for elimination of toxic Cd and dye coupled with ethanol production in desirable lignocelluloses. Carbohydrate Polymers, 2022, 286, 119298. doi: 10.1016/j.carbpol.2022.119298
	Zhai R, Hu J G, Saddler J N. The inhibition of hemicellulosic sugars on cellulose hydrolysis are highly dependant on the cellulase productive binding, processivity, and substrate surface charges. Bioresource Technology, 2018, 258, 79- 87. doi: 10.1016/j.biortech.2017.12.006
	Zhang G F, Wang L Q, Li X K, et al. Distinctively altered lignin biosynthesis by site-modification of OsCAD2 for enhanced biomass saccharification in rice. GCB Bioenergy, 2021, 13 (2): 305- 319. doi: 10.1111/gcbb.12772
	Zhang S X, Sheng H C, Ma Y, et al. Mutation of CESA1 phosphorylation site influences pectin synthesis and methylesterification with a role in seed development. Journal of Plant Physiology, 2022, 270, 153631. doi: 10.1016/j.jplph.2022.153631
	Zhang T, Tang H S, Vavylonis D, et al. Disentangling loosening from softening: insights into primary cell wall structure. The Plant Journal:for Cell and Molecular Biology, 2019, 100 (6): 1101- 1117. doi: 10.1111/tpj.14519
	Zhao J Y, Chen H Z. Stimulation of cellulases by small phenolic compounds in pretreated stover. Journal of Agricultural and Food Chemistry, 2014, 62 (14): 3223- 3229. doi: 10.1021/jf405046m
	Zhao X B, Zhang L H, Liu D H. 2012. Biomass recalcitrance. Part I: the chemical compositions and physical structures affecting the enzymatic hydrolysis of lignocellulose. Biofuels, Bioproducts and Biorefining, 6(4): 465-482.
	Zhao X X, Meng X Z, Ragauskas A J, et al. Unlocking the secret of lignin-enzyme interactions: recent advances in developing state-of-the-art analytical techniques. Biotechnology Advances, 2022, 54, 107830. doi: 10.1016/j.biotechadv.2021.107830
	Zhao Y Y, Man Y, Wen J L, et al. Advances in imaging plant cell walls. Trends in Plant Science, 2019, 24 (9): 867- 878. doi: 10.1016/j.tplants.2019.05.009

类别 Category	生物质 Plant species	纤维素 Cellulose (%）	半纤维素 Hemicellulose (%)	木质素 Lignin (%)	参考文献 Reference
草本Herbaceous plant	稻Oryza sativa	35.80	21.50	11.90	Mankar et al., 2021
	小麦Triticum aestivum	31.50	22.60	5.30	Holopainen-Mantila et al., 2013
	芦苇Phragmites australis	77.98	23.56	20.01	胡惠仁等，2000
	荻 Miscanthus sacchariflorus	77.42	22.42	22.71	胡惠仁等，2000
	芒Miscanthus sinensis	49.00	32.00	28.00	Hu et al., 2017
	芦竹Arundo donax	28.02	13.89	20.29	Proietti et al., 2017
	甘蔗Saccharum	38.20	27.10	20.20	Karthikeyan et al., 2013
	柳枝稷Panicum virgatum	32.87	29.12	19.03	彭霄鹏等，2019
	毛竹 Phyllostachys edulis	45.30	17.00	27.00	Ling et al., 2022
	龟甲竹Phyllostachys heterocycla	40.40	24.30	31.60	He et al., 2022
	玉米Zea mays	40.00	25.00	17.00	Machado et al., 2016
	高粱Sorghum bicolor	37.00	33.00	20.00	Wei et al., 2021
灌木 Shrub	柠条Caragana korshinskii	42.70	22.24	22.24	Liu et al., 2022
	灌木柳Salix purpurea	48.89	21.08	24.08	Gouker et al., 2021
	藜蒿Grewia flavescens	58.46	15.32	12.51	Tiwari et al., 2022
	岩蔷薇Cistus salvifolius	11.90	10.20	7.91	Bruno-Soares et al., 2011
针叶材Coniferous trees	欧洲云杉Picea abies	44.50	18.30	26.80	Pielhop et al., 2017
	欧洲赤松Pinus sylvestris	55.90	14.18	22.49	Lin et al., 2022
	冷杉Abies pectinata	46.78	19.28	22.49	Lin et al., 2022
阔叶材Broadleaf trees	欧洲水青冈Fagus sylvatica	44.87	33.53	16.75	Lin et al., 2022
	桉Eucalyptus	49.13	17.45	26.17	Rigual et al., 2018
	欧洲黑松Pinus nigra	29.56	41.92	19.99	Gupta et al., 2014
	沼生栎 Quercus palustris	40.40	35.90	24.10	Isikgor et al., 2015

[1]	陈介南王义强何钢章怀云周再魁. 木质纤维生产燃料乙醇的生物转化技术[J]. 林业科学, 2007, 43(5): 99-105.
[2]	张晓燕;赵广杰刘志军;. 木质生物质的生物分解及生物转化研究进展[J]. 林业科学, 2006, 42(3): 85-93.