СОВРЕМЕННЫЕ ДОСТИЖЕНИЯ В ПРИМЕНЕНИИ ЦЕЛЛЮЛОЗОЛИТИЧЕСКИХ МИКРООРГАНИЗМОВ ДЛЯ ПЕРЕРАБОТКИ ЛИГНОЦЕЛЛЮЛОЗНОЙ БИОМАССЫ

Лигноцеллюлозная биомасса (ЛЦБ) представляет собой важный ресурс для получения биотоплива и других высокоценных продуктов. Основными компонентами ЛЦБ являются целлюлоза и лигнин, которые сложно разлагаются. Исследования последних лет сосредоточились на применении целлюлозолитических микроорганизмов, таких как бактерии Bacillus subtilis, грибы Trichoderma reesei и Penicillium oxalicum, для эффективного расщепления этих компонентов. В статье представлен обзор современных достижений, включая применение глубоких эвтектических растворителей (DES), что показало улучшение доступности целлюлозы и повышение выхода сахаров до 80%. Применение агропромышленных отходов, таких как рисовая солома, в качестве субстрата для производства ферментов, способствует снижению стоимости производства и поддерживает циркулярную биоэкономику.
Описаны стратегии повышения эффективности ферментативной переработки,  включая разработку мультиферментных комплексов и генетически модифицированных штаммов микроорганизмов. Например, использование многофункциональных ферментов из Clostridium cellulosi позволило увеличить выход сахаров на 30%. Рассмотрены возможности интеграции ко-культур грибов и гидротермальных методов обработки, которые обеспечивают синергетический эффект в переработке ЛЦБ.
Особое внимание уделено ключевым ферментам, таким как целлюлазы, гемицеллюлазы и лигниназы, и их роли в расщеплении растительных полимеров.
Обсуждаются перспективы дальнейших исследований, включая разработку термостабильных ферментов, внедрение экологически чистых методов обработки и интеграцию новых технологий в промышленные биорефинерии.

1. Deep eutectic solvents as promising pretreatment agents for sustainable lignocellulosic biorefineries: a review / V. Sharma et al // Bioresour. Technol. – 2022. – Vol. 360. – P. 127631. https://doi.org/10.1016/j.biortech.2022.127631.

2. Agro-industrial food waste as a low-cost substrate for sustainable production of industrial enzymes: a critical review / V. Sharma et al // Catalysts. – 2022. – Vol. 12. – P. 1373. https://doi.org/10.3390/catal12111373.

3. Agricultural waste biorefinery development towards circular bioeconomy / M.K. Awasthi et al // Renew. Sustain. Energy Rev. – 2022. – Vol. 158. – P. 112122. https://doi.org/10.1016/j.rser.2022.112122.

4. Rojas L.F. Agro-industrial waste enzymes: perspectives in circular economy / L.F. Rojas, P. Zapata, L. Ruiz-Tirado // Curr. Opin. Green Sustain. Chem. – 2022. – Vol. 34. – P. 100585. https://doi.org/10.1016/j.cogsc.2021.100585.

5. Lignocellulosic biomass valorization for bioethanol production: a circular bioeconomy approach / А. Devi et al // Bioenergy Res. – 2022. https://doi.org/10.1007/s12155-022-10401-9.

6. Devi M.M. Rice straw: a major renewable lignocellulosic biomass for value-added carbonaceous materials. / M.M. Devi, N. Aggarwal, S. Saravanamurugan // Curr. Green Chem. – 2020. – Vol. 7. – P. 290-303. https://doi.org/10.2174/2213346106666191127120259.

7. Review on neoteric biorefinery systems from detritus lignocellulosic biomass: a profitable approach / N. Kaur et al // J. Clean. Prod. – 2020. – Vol. 256. – P. 120607. https://doi.org/10.1016/j.jclepro.2020.120607.

8. Saini S. Fungal lignocellulolytic enzymes and lignocellulose: a critical review on their contribution to multiproduct biorefinery and global biofuel research / S. Saini, K.K. Sharma // Int. J. Biol. Macromol. – 2021. – Vol. 193. – P. 2304-2319. https://doi.org/10.1016/j.ijbiomac.2021.11.063.

9. Role of systematic biology in biorefining of lignocellulosic residues for biofuels and chemicals production / V. Sharma et al // Sustainable Biotechnology: Enzymatic Resources of Renewable Energy. – 2018. – Р. 5-55. https://doi.org/10.1007/978-3-319-95480-6_2.

10. Nargotra P. Cellulase production from Bacillus subtilis SV1 and its application potential for saccharification of ionic liquid pretreated pine needle biomass under one-pot consolidated bioprocess / P. Nargotra, S. Vaid, B.K. Bajaj // Fermentation. – 2016. – Vol. 2. – P. 19. https://doi.org/10.3390/fermentation2040019.

11. Chapman J. Industrial applications of enzymes: recent advances, techniques, and outlooks / J. Chapman, A.E. Ismail, C.Z. Dinu // Catalysts. – 2018. – Vol. 8. – P. 238. https://doi.org/10.3390/catal8060238.

12. The carbohydrate-active enzymes database (CAZy) // V. Lombard et al // Nucleic Acids Res. – 2014. – Vol. 42. – Р. D490-D495. https://doi.org/10.1093/nar/gkt1178.

13. Synergistic cellulose hydrolysis dominated by a multi-modular processive endoglucanase from Clostridium cellulosi / M. Yang et al // Front. Microbiol. – 2016. – Vol. 7. – P. 932. https://doi.org/10.3389/fmicb.2016.00932.

14. Production of a high-efficiency cellulase complex via β-glucosidase engineering in Penicillium oxalicum / G. Yao et al // Biotechnol. Biofuels. – 2016. – Vol. 9. – P. 78. https://doi.org/10.1186/s13068-016-0491-4.

15. Genetic modification: a tool for enhancing beta-glucosidase production for biofuel application / R.R. Singhania et Al // Bioresour. Technol. – 2017. – Vol. 245. – Р. 1352-1361. https://doi.org/10.1016/j.biortech.2017.05.126.

16. Behera S.S. Solid-state fermentation for production of microbial cellulases: recent advances and improvement strategies / S.S. Behera, R.C. Ray // Int. J. Biol. Macromol. – 2016. – Vol. 86. – Р. 656-669. https://doi.org/10.1016/j.ijbiomac.2015.10.090.

17. Designing a cellulolytic enzyme cocktail for the efficient and economical conversion of lignocellulosic biomass to biofuels. Enzym / М. Adsul et al // Microb. Technol. – 2020. – Vol. 133. – P. 109442. https://doi.org/10.1016/j.enzmictec.2019.109442.

18. Nargotra P. Consolidated bioprocessing of surfactant-assisted ionic liquid-pretreated Parthenium hysterophorus biomass for bioethanol production / P. Nargotra, V. Sharma, B.K. Bajaj // Bioresour. Technol. – 2019. – Vol. 289. – Р. 121611. https://doi.org/10.1016/j.biortech.2019.121611.

19. Insights from enzymatic degradation of cellulose and hemicellulose to fermentable sugars: a review / А.А. Houfani et al // Biomass Bioenergy. – 2020. – Vol. 134. – P. 105481. https://doi.org/10.1016/j.biombioe.2020.105481.

20. Enzymatic processing of lignocellulosic biomass: principles, recent advances and perspectives / H. Ostby et al // J. Ind. Microbiol. Biotechnol. – 2020. – Vol. 47. – P. 623-657. https://doi.org/10.1007/s10295-020-02301-8.

21. Iram A. Ideal feedstock and fermentation process improvements for the production of lignocellulolytic enzymes / A. Iram, D. Cekmecelioglu, A. Demirci // Processes. – 2021. – Vol. 9. – P. 38. https://doi.org/10.3390/pr9010038.

22. Bacterial enzymes involved in lignin degradation / G. De Gonzalo et al // J. Biotechnol. – 2016. – Vol. 236. – Р. 110-119. https://doi.org/10.1016/j.jbiotec.2016.08.011.

23. Plácido J. Ligninolytic enzymes: a biotechnological alternative for bioethanol production / J. Plácido, S. Capareda // Bioresour. Bioprocess. – 2015. – Vol. 2. – P. 23. https://doi.org/10.1186/s40643-015-0049-5.

24. Environment friendly pretreatment approaches for the bioconversion of lignocellulosic biomass into biofuels and value-added products / S. Sharma et al // Environments. – 2023. – Vol. 10. – P. 6. https://doi.org/10.3390/environments10010006.

25. Cellulolytic and xylanolytic enzymes from yeasts: properties and industrial applications / М. Sohail et al // Molecules. – 2022. – Vol. 27. https://doi.org/10.3390/molecules27123783.

26. Bioethanol production from lignocellulosic biomass by environment-friendly pretreatment methods: a review / М. Tayyab et al // Appl. Ecol. Environ. Res. – 2017. – Vol. 16. http://dx.doi.org/10.15666/aeer/1601_225249.

27. Optimization of steam explosion parameters for improved biotechnological use of wheat straw / D. Sulzenbacher et al // Biomass Convers. Biorefinery. – 2023. – Vol. 13. – P. 1035-1046. https://doi.org/10.1007/s13399-020-01266-z.

28. Sperandio G.B. Fungal co-cultures in the lignocellulosic biorefinery context: a review / G.B. Sperandio, E.X. Ferreira Filho // Int. Biodeterioration Biodegrad. – 2019. – Vol. 142. – P. 109-123. https://doi.org/10.1016/j.ibiod.2019.05.014.

29. Enhanced hydrolysis of lignocellulosic biomass with doping of a highly thermostable recombinant laccase / R. Rai et al // Int. J. Biol. Macromol. – 2019. – Vol. 137. – P. 232-237. https://doi.org/10.1016/j.ijbiomac.2019.06.221.

30. Recent advances and challenges of inter-disciplinary biomass valorization by integrating hydrothermal and biological techniques / В. Song et al // Renew. Sustain. Energy Rev. – 2021. – Vol. 135. – P. 110370. https://doi.org/10.1016/j.rser.2020.110370.

31. Production of ethanol from municipal solid waste of India and Nepal. In: Ghosh S.K. (Ed.) / B. Thapa et al // Waste Valorisation and Recycling, Springer Singapore, Singapore. – 2019. – Р. 47-58. https://doi.org/10.1007/978-981-13-2784-1_5.

32. Tsegaye B. Alkali delignification and Bacillus sp. BMP01 hydrolysis of rice straw for enhancing biofuel yields / B. Tsegaye, C. Balomajumder, P. Roy // Bull. Natl. Res. Centre. – 2019. – Vol. 43. – P. 136. https://doi.org/10.1186/s42269-019-0175-x.

33. Tsegaye B. Alkali pretreatment of wheat straw followed by microbial hydrolysis for bioethanol production / B. Tsegaye, C. Balomajumder, P. Roy // Environ. Technol. – 2019. – Vol. 40. – Р. 1203-1211. https://doi.org/10.1080/09593330.2017.1418911.

34. Bioprospecting of functional cellulases from metagenome for second-generation biofuel production: a review / R. Tiwari et al // Crit. Rev. Microbiol. – 2018. – Vol. 44. – P. 244-257. – https://doi.org/10.1080/1040841X.2017.1337713.

35. Tushar M. Efficiency analysis of crude versus pure cellulase in industry / M. Tushar, A. Dutta // Biomass Bioenergy. – 2020. – Р. 283-298. https://doi.org/10.1007/978-981-13-8637-4_10.

36. Vasić K., Ž, Knez, Leitgeb M. Bioethanol production by enzymatic hydrolysis from different lignocellulosic sources. Molecules, 2021, vol. 26. DOI: https://doi.org/10.3390/molecules26030753

37. Optimization of alkali pretreatment to enhance rice straw conversion to butanol / А. Valles et al // Biomass Bioenergy. – 2021. – Vol. 150. – P. 106131. https://doi.org/10.1016/j.biombioe.2021.106131.

38. Biodiesel production from lignocellulosic biomass using Yarrowia lipolytica / М. Vasaki et al // Energy Convers. Manage.: X. – 2022. – Vol. 13. – P. 100167. https://doi.org/10.1016/j.ecmx.2021.100167.

39. A comprehensive review on the framework to valorise lignocellulosic biomass as biorefinery feedstocks / Н.Р. Vu et al // Sci. Total Environ. – 2020. – Vol. 743. – P. 140630. https://doi.org/10.1016/j.scitotenv.2020.140630.

40. Lignocellulosic biomass as sustainable feedstock and materials for power generation and energy storage / F. Wang et al // J. Energy Chem. – 2021. – Vol. 57. – P. 247-280. https://doi.org/10.1016/j.jechem.2020.08.060.

41. Solid-state fermentation parameters effect on cellulase production from empty fruit bunch / V. Wonoputri et al // Bull. Chem. React. Eng. Catal. – 2018. – Vol. 13. – P. 553-559. https://doi.org/10.9767/bcrec.13.3.1964.553-559.

42. Xiang J. Cellulase production from Trichoderma reesei RUT C30 induced by continuous feeding of steam-exploded Miscanthus lutarioriparius. / J. Xiang, X. Wang, T. Sang // Ind. Crops Prod. – 2021. – Vol. 160. – P. 113129. https://doi.org/10.1016/j.indcrop.2020.113129.

43. Bacteria-enhanced dilute acid pretreatment of lignocellulosic biomass / X. Yan et al // Bioresour. Technol. – 2017. – Vol. 245. – P. 419-425. https://doi.org/10.1016/j.biortech.2017.08.037.

44. Improving the saccharification efficiency of lignocellulosic biomass using a bio-inspired twostage microreactor system loaded with complex enzymes / А. Xia et al // Green Chem. – 2022. – Vol. 24. – P. 9519-9529. https://doi.org/10.1039/D2GC02965K.

45. Changes of chemical composition and hemicelluloses structure in differently aged bamboo (Neosinocalamus affinis) culms / P.-P. Yue et al // J. Agricult. Food Chem. – 2018. – Vol. 66. – P. 9199-9208. https://doi.org/10.1021/acs.jafc.8b03516.

46. Enzymatic hydrolysis of lignocellulosic biomass using a novel, thermotolerant recombinant xylosidase enzyme from Clostridium clariflavum: a potential addition for biofuel industry / A. Zafar et al // RSC Adv. – 2022. – Vol. 12. – P. 14917-14931. https://doi.org/10.1039/D2RA00304J.

47. Zhang H. An insight to pretreatment, enzyme adsorption and enzymatic hydrolysis of lignocellulosic biomass: experimental and modeling studies / H. Zhang, L. Han, H. Dong // Renew. Sustain. Energy Rev. – 2021. – Vol. 140. – P. 110758. https://doi.org/10.1016/j.rser.2021.110758.

48. Zhang F. Improvement of cellulase production in Trichoderma reesei rut-C30 by overexpression of a novel regulatory gene Trvib-1 / F. Zhang, X. Zhao, F. Bai // Bioresour. Technol. – 2018. – Vol. 247. – Р. 676-683. https://doi.org/10.1016/j.biortech.2017.09.126.

49. Integrated process to produce biohydrogen from wheat straw by enzymatic saccharification and dark fermentation / J. Zhu et al // Int. J. Hydrogen Energy. – 2022. https://doi.org/10.1016/j.ijhydene.2022.05.056.

50. Use of bacteria for improving the lignocellulose biorefinery process: importance of pre-erosion / S. Zhuo et al // Biotechnol. Biofuels. – 2018. – Vol. 11. – P. 146. https://doi.org/10.1186/s13068-018-1146-4.

51. Lignocellulosic biomass pre-treatment using low-cost ionic liquid for bioethanol production: an economically viable method for wheat straw fractionation / Z. Ziaei-Rad et al // Biomass Bioenergy. – 2021. – Vol. 151. – P. 106140. https://doi.org/10.1016/j.biombioe.2021.106140.

52. Bashir N. Enzyme immobilization and its applications in food processing: a review / N. Bashir, M. Sood, J.D. Bandral // Int. J. Chem. Stud. – 2020. – Vol. 8. – P. 254-261. https://doi.org/10.22271/chemi.2020.v8.i2d.8779.

53. Jerusalem artichoke (Helianthus tuberosus L.): a versatile and sustainable crop for renewable energy production in Europe / F. Rossini et al // Agronomy. – 2019. – Vol. 9. – P. 528. https://doi.org/10.3390/agronomy9090528.

54. Biotransformation of lignocellulosic biomass to value-added bioproducts: insights into biosaccharification strategies and potential topics in catalysis / М. Jahangeer et al // Topics in Catalysis. – 2024. https://doi.org/10.1007/s11244-024-01941-9.

55. Xylitol: production strategies with emphasis on biotechnological approach, scale up, and market trends / S. Mathur et al // Sustainable Chemistry and Pharmacy. – 2023. – Vol. 35. – P. 101203. https://doi.org/10.1016/j.scp.2023.101203.

56. Utilization of agricultural wastes for co-production of xylitol, ethanol, and phenylacetylcarbinol: a review / J. Feng et al // Bioresource Technology. – 2024. – Vol. 392. – P. 129926. https://doi.org/10.1016/j.biortech.2023.129926.

57. Xylitol: bioproduction and applications-a review / D. Umai et al // Frontiers in Sustainability. – 2022. – Vol. 3. – P. 826190. https://doi.org/10.3389/frsus.2022.826190.

58. Biological pretreatment of lignocellulosic biomass: an environment-benign and sustainable approach for conversion of solid waste into value-added products / R.C. Kuhad et al // Critical Reviews in Environmental Science and Technology. – 2023. – Vol. 54. – № 10. https://doi.org/10.1080/10643389.2023.2277670.

59. Получение биогаза отходов сельского хозяйства / Б. Бахтияр и др. // Вестник КАЗАТК. Энергетика. – 2023. – Том 128, № 5. https://doi.org/10.52167/1609-1817-2023-128-5-399-409.