林业科学 ›› 2026, Vol. 62 ›› Issue (3): 1-12.doi: 10.11707/j.1001-7488.LYKX20250058
收稿日期:2025-02-05
修回日期:2025-05-08
出版日期:2026-03-15
发布日期:2026-03-12
通讯作者:
马星霞
E-mail:mxxyln@139.com
基金资助:
Jiayue Zhao1,Zhijie Zong1,Xiyuan Peng1,Zhiqiang Li1,Xingxia Ma2,*(
)
Received:2025-02-05
Revised:2025-05-08
Online:2026-03-15
Published:2026-03-12
Contact:
Xingxia Ma
E-mail:mxxyln@139.com
摘要:
木质素是植物细胞壁的重要组成部分,与纤维素和半纤维素共同构成陆地生态系统中最大的可再生有机碳库。因复杂的分支结构、异质性和多种键型组成,木质素降解往往较为困难,这一特性不仅制约木质素的高值化利用,还在很大程度上限制木质纤维素的高效转化。生物降解是一种绿色、低碳、环保的处理方法,在木质素转化利用中展现出巨大的应用潜力和广阔的发展前景,深入研究木质素的生物降解机制,不仅有助于为生物质转化利用和可再生化学品生产提供新的思路,而且有利于增进对全球碳封存的理解,从而更好应对当前的气候变化和能源危机。真菌作为地球上最主要的分解者,进化出最全面的酶系统用于木质素降解。然而,目前关于真菌降解木质素的研究仍存在诸多空白,其酶系统对木质素的降解机制尚未得到完全阐明。近年来一些研究取得了重要进展,如软腐菌Parascedosporium putredinis NO1分泌一种新型的、不需要辅助因子的木质素氧化酶,可打破木质素结构中的β-醚键,并释放三嗪;白腐菌Echinodontium taxodii可分泌锰过氧化物酶、漆酶和酯酶,这几种胞外酶协同作用对竹子木质素进行选择性脱除,破坏木质素与木聚糖的交联键,从而显著提高纤维素酶对纤维素的可及性和糖化效率。此外,白腐菌Phanerochaete chrysosporium分泌的木质素过氧化物酶PcLiP03通过静电和疏水作用特异性结合于木质素表面,与纤维素的结合能力较弱,纤维素加入几乎不影响PcLiP03对木质素的作用效果。在分解代谢方面,13C同位素标记技术的应用揭示担子菌Agaricus bisporus在胞外对木质素衍生物和天然木质素进行解聚,并以分解产物为碳源和能源,在胞内同化为多种细胞组分。同时,最新研究首次明确证明,厌氧真菌Neocallimastigomycete californiae可通过非特异性的小分子介导木质素解聚过程,改变木质素单体的组成比例,并引发多种木质素的化学键断裂。综上所述,这些研究成果在一定程度上突破了真菌降解木质素的传统认知,为发展更高效、经济的木质素生物降解技术提供了扎实的理论基础,同时为生物地球化学循环提供了全新的见解。下一步研究应进一步明确真菌降解木质素的酶促机制,并继续挖掘真菌代谢路径在木质素高值化利用中的潜力。
中图分类号:
赵佳玥,宗志洁,彭玺元,李志强,马星霞. 真菌降解木质素的酶及其代谢途径研究进展[J]. 林业科学, 2026, 62(3): 1-12.
Jiayue Zhao,Zhijie Zong,Xiyuan Peng,Zhiqiang Li,Xingxia Ma. Research Progress on Enzymes and Metabolic Pathways Involved in Lignin Biodegradation by Fungi[J]. Scientia Silvae Sinicae, 2026, 62(3): 1-12.
表1
新型木质素降解酶c2092_g1_i1与主要木质素氧化酶的特征对比"
| 特征Characteristics | c2092_g1_i1 | LiPs | MnPs | Laccases |
| 类型 Type | 氧化酶 Oxidase | 氧化酶 Oxidase | 氧化酶 Oxidase | 氧化酶 Oxidase |
| 分子量 Molecular weight | 44.5 kDa | 38~43 kDa | 38~62.5 kDa | 40~140 kDa 真菌漆酶Fungal laccase:40?140 kDa |
| 来源 Source | P. putredinis NO1在含小麦秸秆的 培养基的上清液 P. putredinis NO1 supernatant after incubation on wheat straw | 黄孢原毛平革菌 White-rot fungus Phanerochaete chrysosporium | 黄孢原毛平革菌 White-rot fungus Phanerochaete chrysosporium | 柬埔寨一种漆树科植物 A plant from Anacardiaceae in Cambodia |
| 蛋白家族 Protein family | 与中央酪氨酸酶结构域(PF00264) 同源,具有双核Ⅲ型铜结合位点 Homology to central tyrosinase domain (PF00264) with binuclear type-Ⅲ copper-binding site | Ⅱ类过氧化物酶 (PF00141) 含有血红素 ClassⅡ peroxidases (PF00141) with heme group | Ⅱ类过氧化物酶 (PF00141) 含有血红素和 Mn2+结合位点 ClassⅡ peroxidases (PF00141) with heme group and Mn2+ binding site | 多铜氧化酶 (PF00394) 含有Ⅰ、Ⅱ、Ⅲ型铜 结合位点 Multicopper oxidases (PF00394) with typeⅠ, Ⅱ, Ⅲ copper sites |
| 分布 Distribution | 在子囊菌纲类真菌中发现具 有>50%序列一致性的同源蛋白 Homologous proteins (>50 sequence identity) found in Sordariomycetes class of Ascomycetes | 存在于各种白腐菌中Found in various white-rot fungi | 存在于各种白腐菌中Found in various white-rot fungi | 广泛存在于真菌、植物和细菌中 Widely found in fungi, plants, and bacteria |
| 辅因子 Cofactors | 不依赖辅因子 Cofactor-independent | 一般不需要 Not typically required | Mn3?螯合和扩散 Required for Mn3+ chelation and diffusion | ABTS 或 HBT ABTS or HBT |
| 氧化特性 Oxidative nature | 厌氧环境下失活 Near-total loss of activity under anaerobic conditions | 厌氧条件下有活性 Active in anaerobic conditions | 厌氧条件下有活性 Active in anaerobic conditions | 厌氧条件下有活性 Active in anaerobic conditions but activity and mechanisms differ |
| 最佳pH值 Optimal pH | 10 | 根据来源和特定的 LiP 而异 Depend on the source and specific LiP | 根据来源和特定的 MnP 而异 Depend on the source and specific MnP | 根据来源和特定的 漆酶而异 Depend on the source and specific laccase |
| 最佳温度 Optimal temperature | 60 ℃ | 根据来源和特定的 LiP 而异 Depend on the source and specific LiP | 根据来源和特定的 MnP而异 Depend on the source and specific MnP | 根据来源和特定的 漆酶而异 Depend on the source and specific laccase |
| 木质素 降解特性 Lignin degradation | 具有β-醚酶活性、可从木质素中 释放麦黄酮 Effective tricin release and with β-etherase activity | 对酚类结构活性高 High for breaking non-phenolic bonds (β—O—4) | 对酚醛结构活性高 High for phenolic substrates, indirect oxidation via Mn3+ chelates | 对于酚键降解效果中等;介质参与效果增强 Moderate for phenolic bonds, enhanced by mediators |
| 潜在应用 Potential applications | 不依赖辅因子、适用于高温和碱性条件,可用于木质素改性和 木质纤维素降解 Cofactors-free, high-temperature and alkaline conditions, use in lignin modification and lignocellulose degradation | 降解木质素生产生物 燃料 Potential in lignin degradation for biofuel production | 与LiPs或laccases结合 Effective in combination with LiPs or laccases | 用途广泛,可用于生物修复、制浆漂白、生物传感 Versatile, used in bioremediation, pulp bleaching, and biosensors |
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