РОЛЬ ФЕРМЕНТУ ЦЕЛЮЛАЗИ ЯК ЕКОЛОГІЧНОГО АГЕНТУ У РОЗКЛАДАННІ ЦЕЛЮЛОЗИ У ПРИРОДІ ТА ПРОМИСЛОВОСТІ
DOI:
https://doi.org/10.31861/biosystems2026.01.058Ключові слова:
фермент целюлаза, розкладання целюлози, екологічний агент, промисловість, зелені технологіїАнотація
Целюлаза являє собою складний синергетичний комплекс ферментів, що складається переважно з трьох типів: ендоглюканаз, екзоглюканаз та β-глюкозидаз. Взаємодія цих біокаталізаторів забезпечує ефективне розщеплення складної структури природного полісахариду целюлози на прості моно- та дисахариди, такі як глюкоза та целобіоза. В природі цей ензим синтизується широкою різномантністю мікроорганізмів – незалежно або через симбіотичні стосунки – при чому представики типів Fungi та Bacteria виступають як основні деградатори рослинної біомаси. Цей процес деградації целюлози є фундаментальним для глобального циклу вуглецю, тим самим підтримуючи життя на нашій планеті.
В промисловому контексті, цей фермент вироблять твердофазною та глибинною ферментацією Ефективність виходу ферменту контролюється комбінацією факторів: хімічних (склад середовища, включаючи джерела Карбону та Нітрогену), фізичних (температура, аерація, перемішування), біологічних (вибір штамів мікроорганізмів, мутагенез, синергетична взаємодія). Використання сільськогосподарських відходів як субстрату є ключовою стратегією підвищення економічної доцільності виробництва. Завдяки своїй універсальності целюлаза знаходить широке застосування в багатьох секторах економіки. Фермент є важливим для харчової промисловості та виробництва напоїв (виноробство, хлібопекарство, освітлення соків), целюлозно-паперовому виробництві, текстильному виробництві, складу мийних засобів, фармацевтичної промисловості. Крім того, фермент відіграє важливу роль у виробництві альтернативного палива та в екстракції за участі ферменту біологічно активних сполук з рослинної сировини.
Метою цього дослідження є пошук та систематизація інформації про роль ферменту целюлази в ферментативному гідролізі, підкреслюючи його стійкий підхід як процес відточений еволюцією та успішно адаптований для промислових «зелених технологій.
Посилання
1. Alamsjah, F., Aswan, D. R., Agustien, A. (2024). pH and Temperature Optimization of Several Bacterial Isolates from Mangrove Waters in the Mandeh Area to Produce Cellulase Enzyme. OnLine Journal of Biological Sciences, Vol. 24(3), pp. 367–373. https://doi.org/10.3844/ojbsci.2024.367.373
2. Aldred, E. (2009). Carbohydrates. In Pharmacology, pp. 63–72. Elsevier, Amsterdam, Netherlands. https://doi.org/10.1016/b978-0-443-06898-0.00009-8
3. Al-Rousan, W. M., Al-Marazeeq, K. M., Abdullah, M. A., Khalaileh, N. I. A., Angor, M. M., Ajo, R. Y. (2021). Use of enzymatic preparations to improve the productivity and quality of olive oil. Jordan Journal of Agricultural Sciences, Vol. 17(4), pp. 455–469. https://doi.org/10.35516/jjas.v17i4.97
4. Alves, V. D., Fontes, C. M. G. A., Bule, P. (2020). Cellulosomes: highly efficient cellulolytic complexes. Sub-cellular Biochemistry/Subcellular Biochemistry, 96, 323–354. https://doi.org/10.1007/978-3-030-58971-4_9
5. Aziz, T., Farid, A., Haq, F., Kiran, M., Ullah, A., Zhang, K., Li, C., Ghazanfar, S., Sun, H., Ullah, R., Ali, A., Muzammal, M., Shah, M., Akhtar, N., Selim, S., Hagagy, N., Samy, M., Jaouni, S. K. A. (2022). A review on the modification of cellulose and its applications. Polymers, Vol. 14(15), 3206. https://doi.org/10.3390/polym14153206
6. Barati, B., Amiri, I. S. (2015). In silico engineering of disulphide bonds to produce stable cellulase. In SpringerBriefs in applied sciences and technology, Springer Singapore, Singapore. https://doi.org/10.1007/978-981-287-432-0
7. Bautista-Cruz, A., Aquino-Bolaños, T., Hernández-Canseco, J., Quiñones-Aguilar, E. E. (2024). Cellulolytic Aerobic Bacteria Isolated from Agricultural and Forest Soils: An Overview. Biology, Vol. 13(2), 102. https://doi.org/10.3390/biology13020102
8. Beimforde, C., Feldberg, K., Nylinder, S., Rikkinen, J., Tuovila, H., Dörfelt, H., Gube, M., Jackson, D. J., Reitner, J., Seyfullah, L. J., Schmidt, A. R. (2014). Estimating the Phanerozoic history of the Ascomycota lineages: Combining fossil and molecular data. Molecular Phylogenetics and Evolution, Vol. 78, pp. 386–398. https://doi.org/10.1016/j.ympev.2014.04.024
9. Bhardwaj, N., Kumar, B., Agrawal, K., Verma, P. (2021). Current perspective on production and applications of microbial cellulases: a review. Bioresources and Bioprocessing, Vol. 8(1), 95. https://doi.org/10.1186/s40643-021-00447-6
10. Bhati, N., Shreya, Sharma, A. K. (2020). Cost‐effective cellulase production, improvement strategies, and future challenges. Journal of Food Process Engineering, Vol. 44(2). https://doi.org/10.1111/jfpe.13623
11. Biotech, E. (Cellulase). Fermenty ENZIM Biotech. https://ferment.enzim.biz/tselulaza.html [In Ukrainian]
12. Biswas, S., Saber, M. A., Tripty, I. A., Karim, M. A., Islam, M. A., Hasan, M. S., Alam, A. S. M. R. U., Jahid, M. I. K., Hasan, M. N. (2020). Molecular characterization of cellulolytic (endo- and exoglucanase) bacteria from the largest mangrove forest (Sundarbans), Bangladesh. Annals of Microbiology, Vol. 70(1). https://doi.org/10.1186/s13213-020-01606-4
13. Bridžiuvienė, D., Raudonienė, V., Švedienė, J., Paškevičius, A., Baužienė, I., Vaitonis, G., Šlepetienė, A., Šlepetys, J., Kačergius, A. (2022). Impact of Soil Chemical Properties on the Growth Promotion Ability of Trichoderma ghanense, T. tomentosum and Their Complex on Rye in Different Land-Use Systems. Journal of Fungi, Vol. 8(1), 85. https://doi.org/10.3390/jof8010085
14. Buffo, M. M., Esperança, M. N., Farinas, C. S., Badino, A. C. (2020). Relation between pellet fragmentation kinetics and cellulolytic enzymes production by Aspergillus niger in conventional bioreactor with different impellers. Enzyme and Microbial Technology, Vol. 139, 109587. https://doi.org/10.1016/j.enzmictec.2020.109587
15. Buffo, M., Corrêa, L., Esperança, M., Cruz, A., Farinas, C., Badino, A. (2016). Influence of dual-impeller type and configuration on oxygen transfer, power consumption, and shear rate in a stirred tank bioreactor. Biochemical Engineering Journal, Vol. 114, pp. 130–139. https://doi.org/10.1016/j.bej.2016.07.003
16. Cabrera, M. Á., Blamey, J. M. (2018). Biotechnological applications of archaeal enzymes from extreme environments. Biological Research, Vol. 51(1), 37. https://doi.org/10.1186/s40659-018-0186-3
17. Cellulase - Worthington Enzyme Manual. Worthington Biochemical. https://www.worthington-biochem.com/products/cellulase/manual
18. Chatterjee, S., Sharma, S., Prasad, R. K., Datta, S., Dubey, D., Meghvansi, M. K., Vairale, M. G., Veer, V. (2016). Cellulase Enzyme based Biodegradation of Cellulosic Materials: An Overview. South Asian Journal of Experimental Biology, Vol. 5(6), pp. 271–282. https://doi.org/10.38150/sajeb.5(6).p271-282
19. Chauhan, J., Shukla, R., Bishoyi, A. K., Goyal, S., Sanghvi, G. (2023). Investigation of physical, nutritional and sensory properties of wheat bread treated with purified thermostable cellulase and alpha amylase. Cogent Food & Agriculture, Vol. 9(1). https://doi.org/10.1080/23311932.2023.2261839
20. Chen, B., Yang, Y., Chen, L., Jiang, L., Hong, Y., Zhu, J., Liu, J., Xu, D., Kuang, K., He, Z. (2023). Microclimate along an elevational gradient controls foliar litter cellulose and lignin degradation in a subtropical forest. Frontiers in Forests and Global Change, Vol. 6. https://doi.org/10.3389/ffgc.2023.1134598
21. Chen, H. (2014). Biotechnology of lignocellulose, 1th ed., Chemical Industry Press, Beijing, China. https://doi.org/10.1007/978-94-007-6898-7
22. Chen, L., Wang, C., Su, J. (2023). Understanding the Effect of Different Glucose Concentrations in the Oligotrophic Bacterium Bacillus subtilis BS-G1 through Transcriptomics Analysis. Microorganisms, Vol. 11(10), 2401. https://doi.org/10.3390/microorganisms11102401
23. Chen, Y., Liu, Y., Zhang, J., Yang, W., Deng, C., He, R. (2019). Cumulative cellulolytic enzyme activities and initial litter quality in prediction of cellulose degradation in an alpine meadow of the eastern Tibetan Plateau. Journal of Plant Ecology, Vol. 13(1), pp. 51–58. https://doi.org/10.1093/jpe/rtz044
24. Daniel, G. (2016). Fungal degradation of wood cell walls. In Secondary Xylem Biology, pp. 131–167, Academic press, New York, USA. https://doi.org/10.1016/b978-0-12-802185-9.00008-5
25. Datta, R. (2024). Enzymatic degradation of cellulose in soil: A review. Heliyon, Vol. 10(1), e24022. https://doi.org/10.1016/j.heliyon.2024.e24022
26. De Almeida, M. N., Falkoski, D. L., Guimarães, V. M., De Rezende, S. T. (2019). Study of gamba grass as carbon source for cellulase production by Fusarium verticillioides and its application on sugarcane bagasse saccharification. Industrial Crops and Products, Vol. 133, 33–43. https://doi.org/10.1016/j.indcrop.2019.03.008
27. De Oliveira Costa, B., Nahas, E. (2012). Growth and enzymatic responses of phytopathogenic fungi to glucose in culture media and soil. Brazilian Journal of Microbiology, Vol. 43(1), pp. 332–340. https://doi.org/10.1590/s1517-83822012000100039
28. Dehghani, F., Reitz, T., Schlüter, S., Kästner, M., Blagodatskaya, E. (2025). Decoupling of heat and CO2 release during decomposition of cellulose and its building blocks in soil. Soil Biology and Biochemistry, Vol. 206, 109801. https://doi.org/10.1016/j.soilbio.2025.109801
29. Doan, C. T., Tran, T. N., Pham, T. P., Tran, T. T. T., Truong, B. P., Nguyen, T. T., Nguyen, M., Bui, T. Q. H., Nguyen, A. D., Wang, S. (2024). Production, Purification, and Characterization of a Cellulase from Paenibacillus elgii. Polymers, Vol. 16(14), 2037. https://doi.org/10.3390/polym16142037
30. Dodge, T. (2010). Production of industrial enzymes. In Enzymes in Food Technology. 2nd ed., Vol. 3, pp. 44–58, Blackwell Publishing Ltd, New Jersey, USA. https://doi.org/10.1002/9781444309935.ch3
31. Dutt, D., Kumar, A. (2014). Optimization of cellulase production under solid-state fermentation by aspergillus flavus (at-2) and Aspergillus niger (at-3) and its impact on stickies and ink particle size of sorted office paper. Cellulose chemistry and technology, Vol. 48(3–4), 285–298.
32. Ejaz, U., Sohail, M., Ghanemi, A. (2021). Cellulases: from bioactivity to a variety of industrial applications. Biomimetics, Vol. 6(3), 44. https://doi.org/10.3390/biomimetics6030044
33. Etale, A., Onyianta, A. J., Turner, S. R., Eichhorn, S. J. (2023). Cellulose: A review of water interactions, applications in composites, and water treatment. Chemical Reviews, Vol. 123(5), pp. 2016–2048. https://doi.org/10.1021/acs.chemrev.2c00477
34. Faheina, G. S., Junior, Sousa, K. A., Zilli, J. E., Vergara, C., Pinto, G. a. S., Santiago-Aguiar, R. S. (2022). Enhanced Cellulase Production by Talaromyces amestolkiae CMIAT055 Using Banana Pseudostem. Waste and Biomass Valorization, Vol. 13(8), pp. 3535–3546. https://doi.org/10.1007/s12649-022-01736-7
35. Foroughi, F., Ghomi, E. R., Dehaghi, F. M., Borayek, R., Ramakrishna, S. (2021). A review on the life cycle assessment of cellulose: from properties to the potential of making it a low carbon material. Materials, Vol. 14(4), 714. https://doi.org/10.3390/ma14040714
36. Goodsell, D. (2023). Cellulases and bioenergy. RCSB Protein Data Bank. https://doi.org/10.2210/rcsb_pdb/mom_2023_5
37. Han, Z., Zhu, H., Cheng, J. (2022). Structure modification and property improvement of plant cellulose: Based on emerging and sustainable nonthermal processing technologies. Food Research International, Vol. 156. https://doi.org/10.1016/j.foodres.2022.111300
38. He, J., Qiu, Y., Ji, X., Liu, X., Qiu, Z., Xu, J., Xia, J. (2021). A novel strategy for producing cellulase from Trichoderma reesei with ultrasound‐assisted fermentation using spent mushroom substrate. Process Biochemistry, Vol. 104, pp. 110–116. https://doi.org/10.1016/j.procbio.2021.03.015
39. Hennessey-Ramos, L., Murillo-Arango, W., Vasco-Correa, J., Astudillo, I. C. P. (2021). Enzymatic Extraction and Characterization of Pectin from Cocoa Pod Husks (Theobroma cacao L.) Using Celluclast® 1.5 L. Molecules, Vol. 26(5), 1473. https://doi.org/10.3390/molecules26051473
40. Hsin, K., Lee, H., Huang, Y., Lin, G., Lin, P., Lin, Y. J., Chen, P. (2025). Lignocellulose degradation in bacteria and fungi: cellulosomes and industrial relevance. Frontiers in Microbiology, Vol. 16, 1583746. https://doi.org/10.3389/fmicb.2025.1583746
41. Huang, X., Fan, W., Wang, S., Xiong, J., Chen, Y., Xie, C. (2024). Highly effective removal of nitrate from saline wastewater by glucose-enhanced sulfur autotrophic system. Journal of Water Process Engineering, Vol. 63, 105439. https://doi.org/10.1016/j.jwpe.2024.105439
42. Huynh, G. H., Van Pham, H., Nguyen, H. V. H. (2023). Effects of enzymatic and ultrasonic-assisted extraction of bioactive compounds from cocoa bean shells. Journal of Food Measurement & Characterization, Vol. 17(5), pp. 4650–4660. https://doi.org/10.1007/s11694-023-01986-6
43. Ilić, N., Milić, M., Beluhan, S., Dimitrijević-Branković, S. (2023). Cellulases: From lignocellulosic biomass to improved production. Energies, Vol. 16(8), 3598. https://doi.org/10.3390/en16083598
44. Intasit, R., Cheirsilp, B., Suyotha, W., Boonsawang, P. (2021). Synergistic production of highly active enzymatic cocktails from lignocellulosic palm wastes by sequential solid state-submerged fermentation and co-cultivation of different filamentous fungi. Biochemical Engineering Journal, Vol. 173, 108086. https://doi.org/10.1016/j.bej.2021.108086
45. Islam, M., Sarkar, P. K., Mohiuddin, A., Suzauddula, M. (2019). Optimization of fermentation condition for cellulase enzyme production from Bacillus sp. Malaysian Journal of Halal Research, Vol. 2(2), pp. 19–24. https://doi.org/10.2478/mjhr-2019-0009
46. Ismaiel, M. M. S., Ahmed, A. E. I., Sobhy, S. (2017). Enhancement of Wheat Cultivars (Triticum aestivum L.) by Cellulase-Treated Plant Wastes. Waste and Biomass Valorization, Vol. 10(6), pp. 1539–1546. https://doi.org/10.1007/s12649-017-0159-8
47. Jain, L., Agrawal, D. (2020). Biofuel cellulases. In Microbial Fermentation and Enzyme Technology, 1st ed., pp. 283–298, CRC Press, Boca Raton, USA. https://doi.org/10.1201/9780429061257-18
48. Jamir, E., Kangabam, R. D., Borah, K., Tamuly, A., Boruah, H. P. D., Silla, Y. (2019). Role of soil microbiome and enzyme activities in plant growth nutrition and ecological restoration of soil health. In Microorganisms for sustainability, pp. 102–103, Springer, Singapore, Singapore. https://doi.org/10.1007/978-981-13-9117-0_5
49. Jayasekara, S., Ratnayake, R. (2019). Microbial cellulases: an Overview and Applications. In Cellulose, Intechopen. https://doi.org/10.5772/intechopen.84531
50. Juturu, V., Wu, J. C. (2014). Microbial cellulases: Engineering, production and applications. Renewable and Sustainable Energy Reviews, Vol. 33, pp.188–203. https://doi.org/10.1016/j.rser.2014.01.077
51. Korsa, G., Konwarh, R., Masi, C., Ayele, A., Haile, S. (2023). Microbial cellulase production and its potential application for textile industries. Annals of Microbiology, Vol. 73(1). https://doi.org/10.1186/s13213-023-01715-w
52. Kracher, D., Ludwig, R. (2016). Cellobiose dehydrogenase: An essential enzyme for lignocellulose degradation in nature – A review / Cellobiosedehydrogenase: Ein essentielles Enzym für den Lignozelluloseabbau in der Natur – Eine Übersicht. Die Bodenkultur Journal of Land Management Food and Environment, Vol. 67(3), pp. 145–163. https://doi.org/10.1515/boku-2016-0013
53. Kulakivska, A., and Konechna, R. (2024). Perspectives of using enzymatically modified raw materials for obtaining bioactive substances. IV International Scientific and Practical Conference «Problems and Achievements of Modern Biotechnology», Kharkiv, Ukraine, pp. 246-249.
54. Kumar, R., Singh, S., Singh, O. V. (2008). Bioconversion of lignocellulosic biomass: biochemical and molecular perspectives. Journal of Industrial Microbiology & Biotechnology, Vol. 35(5), pp. 377–391. https://doi.org/10.1007/s10295-008-0327-8
55. Lenhardt, K. R., Brandt, L., Poll, C., Rennert, T., Kandeler, E. (2023). Release of glucose from dissolved and mineral‐bound organic matter by enzymatic hydrolysis. European Journal of Soil Science, Vol. 74(5). https://doi.org/10.1111/ejss.13421
56. Li, Q., Lei, Y., Hu, G., Lei, Y., Dan, D. (2018). Effects of Tween 80 on the liquid fermentation of Lentinus edodes. Food Science and Biotechnology, Vol. 27(4), pp. 1103–1109. https://doi.org/10.1007/s10068-018-0339-8
57. Li, Q., Wang, S., Wu, G., Tan, Y., Liu, Y., Yuan, C., Geng, S., Liu, Y. (2023). Physiological and biochemical changes in leaf abscission of Cyclocarya paliurus stem segments in vitro. Plant Cell Tissue and Organ Culture (PCTOC), Vol. 155(3), 773–783. https://doi.org/10.1007/s11240-023-02598-0
58. Liu, L., Huang, W., Liu, Y., Li, M. (2021). Diversity of cellulolytic microorganisms and microbial cellulases. International Biodeterioration & Biodegradation, Vol. 163, 105277. https://doi.org/10.1016/j.ibiod.2021.105277
59. Liu, P., Lin, A., Zhang, G., Zhang, J., Chen, Y., Shen, T., Zhao, J., Wei, D., Wang, W. (2019). Enhancement of cellulase production in Trichoderma reesei RUT-C30 by comparative genomic screening. Microbial Cell Factories, Vol. 18(1), 81. https://doi.org/10.1186/s12934-019-1131-z
60. Liu, Z., Li, H., Cui, G., Wei, M., Zou, Z., Ni, H. (2021). Efficient extraction of essential oil from Cinnamomum burmannii leaves using enzymolysis pretreatment and followed by microwave-assisted method. LWT, Vol. 147, 111497. https://doi.org/10.1016/j.lwt.2021.111497
61. Łubek-Nguyen, A., Ziemichód, W., Olech, M. (2022). Application of Enzyme-Assisted extraction for the recovery of natural bioactive compounds for nutraceutical and pharmaceutical applications. Applied Sciences, Vol. 12(7), 3232. https://doi.org/10.3390/app12073232
62. Mäkelä, M. R., Hildén, K. S., Kuuskeri, J. (2020). Fungal Lignin-Modifying peroxidases and H2O2-Producing enzymes. In Encyclopedia of Mycology, pp. 247–259, Elsiver. https://doi.org/10.1016/b978-0-12-809633-8.21127-8
63. Martău, G., Unger, P., Schneider, R., Venus, J., Vodnar, D. C., López-Gómez, J. P. (2021). Integration of solid state and submerged fermentations for the valorization of organic municipal solid waste. Journal of Fungi, Vol. 7(9), 766. https://doi.org/10.3390/jof7090766
64. Mendoza, N. N. G., Rodríguez, S. a. V., Lima, B. L. R. (2020). Improvement of the extraction of carotenoids and capsaicinoids of chili pepper native (Capsicum baccatum), assisted with cellulolytic enzymes. Revista Peruana De Biología, Vol. 27(1), pp. 55–60. https://doi.org/10.15381/rpb.v27i1.17588
65. Metreveli, E., Khardziani, T., Elisashvili, V. (2021). The carbon source controls the secretion and yield of Polysaccharide-Hydrolyzing enzymes of basidiomycetes. Biomolecules, Vol. 11(9), 1341. https://doi.org/10.3390/biom11091341
66. Naher, L., Fatin, S. N., Sheikh, M. a. H., Azeez, L. A., Siddiquee, S., Zain, N. M., Karim, S. M. R. (2021). Cellulase Enzyme Production from Filamentous Fungi Trichoderma reesei and Aspergillus awamori in Submerged Fermentation with Rice Straw. Journal of Fungi, Vol. 7(10), 868. https://doi.org/10.3390/jof7100868
67. New and Future Developments in Microbial Biotechnology and Bioengineering. (2019), 1st ed., Elsevier https://doi.org/10.1016/c2018-0-01234-7
68. Nguyen, S. T. C., Freund, H. L., Kasanjian, J., Berlemont, R. (2018). Function, distribution, and annotation of characterized cellulases, xylanases, and chitinases from CAZy. Applied Microbiology and Biotechnology, Vol. 102(4), p. 1629–1637. https://doi.org/10.1007/s00253-018-8778-y
69. Nie, H., Wang, C., Tian, M., Gao, J. (2023). Exogenous enzyme addition affects litter decomposition by altering the microbial community and litter nutrient content in planted forest. Journal of Plant Ecology, Vol. 16(6). https://doi.org/10.1093/jpe/rtad031
70. Oancea, S., Perju, M. (2020). Influence of enzymatic and ultrasonic extraction on phenolics content and antioxidant activity of Hibiscus Sabdariffa L. flowers. Bulgarian Chemical Communications, Vol. 52, pp. 25-29.
71. Obeng, E. M., Adam, S. N. N., Budiman, C., Ongkudon, C. M., Maas, R., Jose, J. (2017). Lignocellulases: a review of emerging and developing enzymes, systems, and practices. Bioresources and Bioprocessing, Vol. 4(1). https://doi.org/10.1186/s40643-017-0146-8
72. Okonkwo, I. F. (2019). Effect of Metal Ions and Enzyme Inhibitor on the Activity of Cellulase Enzyme of Aspergillus flavus. International Journal of Environment Agriculture and Biotechnology, Vol. 4(3), pp. 727–734. https://doi.org/10.22161/ijeab/4.3.20
73. Olatunji, K. O., Ahmed, N. A., Ogunkunle, O. (2021). Optimization of biogas yield from lignocellulosic materials with different pretreatment methods: a review. Biotechnology for Biofuels, Vol. 14(1), 159. https://doi.org/10.1186/s13068-021-02012-x
74. Onyia, D., Onyeneke, E., Okunwaye, T., Okogbenin, E., Asiriuwa, N., Obibuzor, J., Anemene, H. (2022). The influence of metal ions on cellulolytic activities of fungal isolates from palm biomass. Nigerian Journal of Biotechnology, Vol. 38(2), 67–72. https://doi.org/10.4314/njb.v38i2.7
75. Orengo, C., Michie, A., Jones, S., Jones, D., Swindells, M., Thornton, J. (1997). CATH – a hierarchic classification of protein domain structures. Structure, Vol. 5(8), pp. 1093–1109. https://doi.org/10.1016/s0969-2126(97)00260-8
76. Ozyilmaz, G., Gunay, E. (2022). Clarification of apple, grape and pear juices by co-immobilized amylase, pectinase and cellulase. Food Chemistry, Vol. 398, 133900. https://doi.org/10.1016/j.foodchem.2022.133900
77. Patel, K., Amaresan, N. (2022). Mass multiplication, production cost, and marketing of cellulase. In Industrial Microbiology Based Entrepreneurship, pp. 37-50, Springer, Singapore, Singapore
78. Pham, V. H. T., Kim, J., Shim, J., Chang, S., Chung, W. (2022). Coconut Mesocarp-Based Lignocellulosic Waste as a Substrate for Cellulase Production from High Promising Multienzyme-Producing Bacillus amyloliquefaciens FW2 without Pretreatments. Microorganisms, Vol. 10(2), 327. https://doi.org/10.3390/microorganisms10020327
79. PubChem. Cellulase. PubChem. https://pubchem.ncbi.nlm.nih.gov/compound/440950
80. Schilling, J. S., Ai, J., Blanchette, R. A., Duncan, S. M., Filley, T. R., Tschirner, U. W. (2012). Lignocellulose modifications by brown rot fungi and their effects, as pretreatments, on cellulolysis. Bioresource Technology, Vol. 116, pp. 147–154. https://doi.org/10.1016/j.biortech.2012.04.018
81. Sethi, S., Gupta, S. (2022). Optimization of cultural parameters for cellulase enzyme production from fungi. Biolife, Vol. 2(3), pp. 989-996. https://doi.org/10.5281/zenodo.7224954
82. Seveso, A., Mazurkewich, S., Banerjee, S., Poulsen, J. N., Lo Leggio, L., Larsbrink, J. (2024). Polysaccharide utilization loci from Bacteroidota encode CE15 enzymes with possible roles in cleaving pectin-lignin bonds. Applied and Environmental Microbiology, Vol. 90(1), e0176823. https://doi.org/10.1128/aem.01768-23
83. Shirkavand, E., Baroutian, S., Gapes, D. J., Young, B. R. (2015). Combination of fungal and physicochemical processes for lignocellulosic biomass pretreatment – A review. Renewable and Sustainable Energy Reviews, Vol. 54, pp. 217–234. https://doi.org/10.1016/j.rser.2015.10.003
84. Shrivastava, S. (2020). Industrial applications of glycoside hydrolases, Springer Singapore, Singapore. https://doi.org/10.1007/978-981-15-4767-6
85. Silva, J. C. R., Salgado, J. C. S., Vici, A. C., Ward, R. J., Polizeli, M. L. T. M., Guimarães, L. H. S., Furriel, R. P. M., Jorge, J. A. (2019). A novel Trichoderma reesei mutant RP698 with enhanced cellulase production. Brazilian Journal of Microbiology, Vol. 51(2), pp. 537–545. https://doi.org/10.1007/s42770-019-00167-2
86. Sobucki, L., Ramos, R. F., Meireles, L. A., Antoniolli, Z. I., Jacques, R. J. S. (2021). Contribution of enzymes to soil quality and the evolution of research in Brazil. Revista Brasileira De Ciência Do Solo, Vol. 45. https://doi.org/10.36783/18069657rbcs20210109
87. Suleiman, M., Krüger, A., Antranikian, G. (2020). Biomass-degrading glycoside hydrolases of archaeal origin. Biotechnology for Biofuels, Vol. 13(1), 153. https://doi.org/10.1186/s13068-020-01792-y
88. Sutaoney, P., Rai, S. N., Sinha, S., Choudhary, R., Gupta, A., Singh, S. K., Banerjee, P. (2024). Current perspective in research and industrial applications of microbial cellulases. International Journal of Biological Macromolecules, Vol. 264(Pt 1), 130639. https://doi.org/10.1016/j.ijbiomac.2024.130639
89. THE 17 GOALS. Sustainable Development. (2026). https://sdgs.un.org/goals
90. Thoresen, M., Malgas, S., Mafa, M., Pletschke, B. (2021). Revisiting the phenomenon of cellulase action: Not all endo- and Exo-Cellulase interactions are synergistic. Catalysts, Vol. 11(2), 170. https://doi.org/10.3390/catal11020170
91. Tuomela, M., Hatakka, A. (2011). Oxidative fungal enzymes for bioremediation. In Comprehensive Biotechnology, 2nd ed., pp. 183–196, Academic press, New York, USA. https://doi.org/10.1016/b978-0-08-088504-9.00370-6
92. Verma, N., Kumar, V. (2020). Utilization of bottle gourd vegetable peel waste biomass in cellulase production by Trichoderma reesei and Neurospora crassa. Biomass Conversion and Biorefinery, Vol. 12(4), 1105–1114. https://doi.org/10.1007/s13399-020-00727-9
93. Vieira, M. M., Kadoguchi, E., Segato, F., Da Silva, S. S., Chandel, A. K. (2020). Production of cellulases by Aureobasidium pullulans LB83: optimization, characterization, and hydrolytic potential for the production of cellulosic sugars. Preparative Biochemistry & Biotechnology, Vol. 51(2), pp. 153–163. https://doi.org/10.1080/10826068.2020.1799393
94. Weerasinghe, W. M. L. I., Madusanka, D. a. T., Manage, P. M. (2021). Isolation and identification of cellulase producing and sugar fermenting bacteria for Second-Generation bioethanol production. International Journal of Renewable Energy Development, Vol. 10(4), pp. 699–711. https://doi.org/10.14710/ijred.2021.35527
95. Wilson, D. (2009). Cellulases. In Elsevier eBooks, 3rd ed., pp. 252–258, Academic Press, New York, USA. https://doi.org/10.1016/b978-012373944-5.00138-3
96. Yazıcı, S. Ö., Özmen, I. (2020). Optimization for coproduction of protease and cellulase from Bacillus subtilis M-11 by the Box–Behnken design and their detergent compatibility. Brazilian Journal of Chemical Engineering, Vol. 37(1), pp. 49–59. https://doi.org/10.1007/s43153-020-00025-x
97. Zhang, W., Zhao, S., Zhang, L., Xing, Y., Jia, W. (2020). Changes in the cell wall during fruit development and ripening in Fragaria vesca. Plant Physiology and Biochemistry, Vol. 154, 54–65. https://doi.org/10.1016/j.plaphy.2020.05.028
