堆肥微生物群落演替及木质素降解功能微生物强化堆肥机理研究
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摘要
堆肥技术作为一项经济有效的有机固体废物处理与资源化利用技术,是备受关注的重要研究课题之一。但传统堆肥法常因含难降解有机物木质素而堆制效果不佳,且因成品中重金属的潜在威胁而应用受限,阻碍了堆肥资源化技术的发展,故改进堆肥技术以提高堆肥效率、降低重金属毒性非常重要。由于堆肥过程中木质素等有机物的降解主要依赖于微生物群落的共同作用,了解微生物群落演替的基本信息,是发展堆肥新方法与理论的重要基础。因此,以微生物群落演替研究为依据,针对传统堆肥法存在的木质素难降解、重金属污染需控制这两个问题,开发能同时实现高效堆肥和降低成品重金属毒性的堆肥新方法与机理,对推动堆肥技术大跨度发展,实现高效快速堆肥处理具有重要的理论价值与现实意义。
本文探讨了堆肥过程中微生物群落的演替规律,系统地研究了高效木质素降解功能微生物与其胞外酶的行为机制及对堆肥微生物群落结构与活性、木质素降解、腐殖质形成的作用机理,并考察强化堆肥法对堆肥重金属毒性的影响,发展了新型高效的基于木质素降解功能微生物、复合酶的强化堆肥技术与理论。本文的具体研究工作及成果包括以下4个部分内容：
第1部分为堆肥过程微生物群落结构变化与木质纤维素降解的相关性研究。(1)采用醌指纹法研究了堆肥过程中微生物群落的演替。发现微生物群落结构发生了明显改变。高温阶段,Q-9(H2)、Q-10(H2)表征的真菌是半纤维素、纤维素的主要降解者,MK-7、Q-10分别表征的细菌、嗜热真菌是主要木质素降解者,而Q-9表征的黑曲霉、青霉等嗜温木质素降解真菌量少,降解作用有限。降温阶段,Q-9和长链甲基萘醌增多,其与优势醌MK-7分别表征的真菌、放线菌、细菌协同作用,促使木质素大量降解,并呈较高的纤维素降解率。(2)含MK-9(H2)的放线菌分别与MK-9、MK-9(H6)、MK-10(H6)表征的放线菌有良好共存性,含MK-5(H2)的细菌分别与MK-9、MK-9(H2)表征的放线菌友好共存,互促生长繁殖。限制性片段长度多态性分析显示,细菌在高温期最活跃,堆肥过程细菌群落基因多样性变化小。(3)结合主成分、相关性分析等探讨了木质素降解微生物群落的组成及其与木质纤维素降解的相关性。发现Q-9、MK-7、MK-8(H2)、MK-9、 MK-9(H2)、MK-9(H6)、MK-9(H8)和MK-10(H6)表征的嗜温真菌、细菌和放线菌组成的特定群落是起关键作用的木质素降解群落,与木质素降解正相关,含后7种醌的微生物群落还与纤维素降解正相关。(4)证实了木质素的降解显著影响半纤维素、纤维素降解,指出堆肥过程木质素降解真菌的劣势地位限制了木质素的降解,可通过接种木质素降解真菌促进木质素降解、提高堆肥效率。
第2部分着重研究木质素降解功能微生物、胞外酶强化堆肥技术及机理。(1)针对高效木质素降解功能微生物白腐真菌特性与降解机理展开研究。发现温度和含水率是影响该菌固态发酵产酶和降解能力的最主要因素,确定最佳工艺参数：菌种驯化5d,接种量0.8%,吐温80添加量0.3%,温度37℃,含水率85%。证实该菌能严重破坏木质素结构,使苯环开环,难降解的大分子长键烃断裂成易降解的小分子短键烃。提出了用关联分析思想判别白腐菌产酶过程最显著影响因素,建立了GM(1,1)模型预测日产酶量,发现在最显著影响因素含水率不变时,GM(1,1)模型能快速准确地预测产酶,减少实验工作量。(2)建立了木质素降解功能微生物强化堆肥法处理木质纤维素类废物。证实强化堆肥法可增加微生物活性,提高木质素降解率至43.9%,使有机物降解更彻底。(3)进一步研究强化堆肥过程中的微生物群落演替和腐殖质形成。发现强化堆肥法显著提高了成品腐殖化程度(54.1%)和总微生物量,但降低了群落多样性指数、均匀性指数,使Q-9、长链甲基萘醌分别表征的木质素降解真菌、放线菌增多,尤其木质素降解真菌成为优势菌,Q-8、MK-7分别表征的固氮细菌、木质素降解细菌增多,而其它微生物减少。聚类分析和主成分分析显示,强化堆肥法明显改变了微生物群落结构组成,尤其在16～28d影响最大,大大加强了木质素降解真菌、放线菌在整个堆肥体系中的影响力,利于木质素降解。(4)制备了木质素降解功能复合酶,考察了其在稻草、米糠、蔬菜残余、土壤4种典型堆肥基质表面的传输行为与附着性能,并研究其降解天然木质素机理。发现复合酶在稻草、米糠中传输性能最佳,可传输至基质柱深层；在4种基质表面的附着都属有利吸附,利于酶与基质的有效接触。复合酶酶解木质素时,通过氧化苯甲醇单元、脱甲氧基和甲基、断裂部分碳碳双键、醚键与Ca-Cβ键并氧化Ca侧链成羧酸、芳环开环及芳环取代等一系列反应,导致木质素结构中醇羟基、亚甲基、甲氧基、甲基、醚键、碳碳双键、苯环等减少,香草酸、阿魏酸等酯型结构被分解,木质素二聚体解聚成低分子物质,羰基、取代芳环等增多。(6)研究了功能复合酶强化堆肥法对微生物群落碳源代谢、木质素降解的影响。结果显示,添加复合酶显著增强了总有机质的降解和微生物群落碳源代谢能力。Biolog分析表明,复合酶主要影响了微生物群落对17种碳源的利用,大大提高了丙酮酸甲酯、a-环式糊精、D-甘露醇、D-半乳糖醛酸、衣康酸、L-天冬酰胺的利用,降低了D,L-a-磷酸甘油、L-苏氨酸、甘氨酰-L-谷氨酸、腐胺的利用。复合酶显著促进了木质素降解,可能与其既能催化木质素分解,又能增强微生物对酚类化合物、羧酸类等木质素降解中间产物的代谢有关。
第3部分为木质素降解功能微生物强化堆肥法处理Pb污染基质及过程机理研究。(1)探讨了Pb胁迫下木质素降解功能微生物白腐菌的反应行为和降解能力。结果表明,该菌通过菌丝缠绕更致密及分泌胞外物质2种反应行为来抵御Pb毒性,在含400mg L-1Pb2+的液态培养基中能较好生长。Pb污染固态基质处理中,白腐菌定殖能力与水溶交换态Pb减少率(含量高于8.2mg kg-1时)正相关；不同浓度Pb对纤维素酶活、纤维素降解有抑制作用,而低浓度Pb使木聚糖酶、木质素降解酶活性和半纤维素、木质素降解增强。(2)系统研究了黄孢原毛平革菌处理不同浓度Pb污染木质纤维素废物的整个过程,探讨了它对Pb的钝化机制。发现即使初始Pb浓度为400mg kg-1干基质时,该菌仍能降解43.1%的木质纤维素,并形成腐殖质,大大减少游离态Pb。初始Pb浓度为30mg kg-1时,木质纤维素、有机质降解率高达56.8%、64.0%,木质素降解选择性指数较高。经扫描电镜与能谱分析发现,该菌对Pb的钝化可能主要通过：菌丝表面的吸附与离子交换作用,及该菌代谢物的络合作用。(3)研究了木质素降解功能微生物强化堆肥法处理Pb污染垃圾及机理。发现强化堆肥法能经济有效地处理Pb污染垃圾,堆制过程中有较高的微生物量、CO2释放量,使成品中水溶交换态Pb含量为0%、毒性小。且指出水溶性有机C/N变化不宜作为重金属垃圾堆肥的腐熟度评价指标,可考虑改用木质素、粗纤维余量。(4)研究了木质素降解功能微生物强化堆肥法修复Pb污染土壤及机理。发现由于微生物菌丝体对Pb离子的强吸附作用及堆肥过程中生成腐殖质的络合作用,污染土壤经强化堆肥法修复后,土壤微生物活性和群落碳源代谢能力增强,游离态Pb转为以残留态和铁锰氧化物结合态存在。
第4部分着重开展关键生物酶活性及微生物群落演替与木质素降解的映射模型的构建。(1)构建了木质纤维素废物处理体系中酶活与木质素降解率关系的人工神经网络(ANN)模型,该模型中输入酶活值能准确预测木质素的降解。(2)构建了堆肥过程中微生物群落演替与木质素降解率关系的ANN模型,该模型中输入群落信息能有效预测木质素降解。ANN模型在复杂关系解析中具优越性,为定量分析关键酶、微生物群落对木质素降解的影响提供了技术支持,可高效指导有关复合酶、复合菌剂的开发和应用。
本论文研究揭示了堆肥微生物群落演替及木质素降解功能微生物强化堆肥机理,可为科学认识堆肥反应本质提供微生物群落信息,为微生物选育、复合生物促进剂等相关研究的合理开展提供依据；并为高效快速堆肥新方法的发展奠定基础,以克服传统堆肥法处理效率低、腐熟程度差等不足。
Composting, as an economic and effective technology for solid waste treatment, is one of important research subjects. But the application of traditional composting are limited due to the biodegradative recalcitrance of lignin and the stresss of heavy metal in composting materials. So the improvement of composting efficiency and heavy metal control are very important. Since microbial community are responsible for organic matter degradation during composting, the study on microbial community succession is a basis of composting technology and theory researches. To improve composting efficiency and control heavy metal pollution, exploring a new composting technology and theory based on microbial community study are needed. It would promote the development of composting technology, and be of theoretical and practical value for efficient and rapid composting treatment.
This dissertation discussed microbial community succession, and investigated the effects of ligninolytic microorganisms and enzymes on microbial activities, lignin degradation and humus formation in composting. The control of heavy metal are also considered in this study. According to these researches, we developed a new and efficient composting techonology based on ligninolytic microorganisms or enzymes. This dissertation is composed of four sections.
The first section describes the research on the relationship between microbial community successtion and lignocellulose degradation.(1) Microbial populations and their relationship to bioconversion during composting were studied by quinone profiling. The obvious changes in microbial population were observed. Fungi indicated by Q-9(H2) and Q-10(H2) were considered to be the most important hemicellulose and cellulose-degrading microorganisms during thermophilic stage. MK-7and Q-10, indicative of certain bacteria and fungi respectively, were responsible for lignin degradation at the thermophilic stage, whereas the mesophile ligninolytic fungi (Aspergillus niger, Penicillium, and etc.) indicated by Q-9is limited. The highest lignin degradation ratio and good cellulose degradation were found at the cooling stage and were correlated with Q-9, MK-7and long-chain menaquinones attributed to mesophilic fungi, bacteria and actinomycetes, respectively.(2) MK-9(H2) positively correlated with MK-9MK-9(H6) and MK-10(H6) respectively, which suggests coexistence and possibly even cooperativity among the corresponding microorganisms. Significant positive correlation was also suggested for the bacteria indicated by MK-5(H2) and the actinomycetes containing MK-9or MK-9(H2) as major quinone. RFLP analysis showed there were no obvious changes in genetic diversity of bacterial community during composting, and bacterial community is very active during thermophilic stage.(3) Principal component and correlation analyses showed a significant positive correlation between lignin degradation ratio and the total content of Q-9、MK-7、MK-8(H2)、MK-9、MK-9(H2)、MK-9(H6)、MK-9(H8)和MK-10(H6), which indicated that the microbial community consisting of mesophilic fungi, bacteria and actinomycetes played a key role in lignin degradation. And a positive correlation between the latter7quinones and cellulose degradation was found.(4) Lignin degradation significantly affected hemicellulose and cellulose degradation. The inferior position of ligninolytic fungi in composting might be responsible for slow degradation of lignin. It is expected that inoculating ligninolytic fungi would accelerate lignin degradation and promote composting efficiency.
The second section focuses on the study of mechanisms of the composting technology by adding ligninolytic microorganism or enzymes.(1) The characteristics and degradation ability of ligninolytic fungus (white-rot fungus) were studied. Temperature and substrate moisture were confirmed to be the main effect factors. The optimal treament conditions are:5days of fungi incubation,0.8%of inoculum,0.3%of Tween80addition,37℃and85%of water content of substrate. This fungus could destroy lignin structure by the cleavage of aromatic ring and the transformation of long-bond hydrocarbons of giant molecule to the short-bond. The study also applied relational analysis to the enzyme production process of white-rot fungi, and constructed GM(1,1) model which could be used to predicted enzyme production to reduce experimental quantity.(2) Composting of lignocellulosic waste by ligninolytic fungi was studied. By this composting method, microbial activities were enhanced, and lignin degradation ratio reached43.9%, and more completed degradation of organic matter was also found.(3) The higher humification degree (54.1%) and total microbial biomass were observed during the composting process with the inocula of ligninolytic fungi than those in tradition composting. While the low diversity and evenness of microbial community were found. It might be because the lignin-degrading fungi and actinomycetes respectively indicated by Q-9and long-chain menaquinones increased, and bacteria with nitrogen fixation and lignin-degrading bacteria indicated by Q-8and MK-7increased, whereas other microorganisms decreased. Composting method with fungi inocula changed microbial community structure obviously, especially from day16to28, enhanced the effect of lignin-degrading fungi and actinomycetes.(4) The transfer and adsorption behaviours and degradation mechanisms of ligninolytic enzymes (LiP-MnP) were studied. The transfer abilities of ligninolytic enzymes on straw and chaff are better than vegetable residues and soil. The adsorption on these four substrates belonged to favourable adsorption, which ensured the effective contact between enzymes and substrates. Degradation mechanisms of ligninolytic enzymes included:benzyl alcohol unit oxidation, methoxy and methyl group removal, C=C and ether bond and Ca-Cβ cleavage, and aromatic ring cleavage and replacement. Therefore, alcohol hydroxyl,-CH2,-OCH3,-CH3, ether bond, C=C, and aromatic ring in lignin decreased. Vanillic acid and ferulic acid unit in lignin were decomposed, and lignin dimers were degraded into low-weight matters.(6) Carbon source utilization and lignin degradation by microbial community were studied during composting with LiP-MnP enzymes. Addition of LiP-MnP enzymes significantly promoted organic matter degradation and carbon source utilization. It was found by Biolog analysis that the enzymes mainly affected the utilization of17carbon source. The utilization of pyruvic acid methyl ester, a-cyclodextrin, D-mannitol, D-galacturonic acid, itaconic acid and L-asparagine were enhanced, whereas the utilization of D,L-a-glycerol phosphate, L-threonine, glycyl-L-glutamic acid and putrescine were weakened. Lignin degradation was significantly promoted by LiP-MnP enzymes. There might be two reasons:the enzymes catalyzed lignin degradation, and enhanced the utilization of lignin degradation products such as phenolic compound and carboxylic acid.
The third section describes the research on composting of Pb-polluted substrates by inocula of ligninolytic fungi.(1) Mycelial growth and the degradation of lignocellulosic waste by ligninolytic fungus Phanerochaete chrysosporium under lead stress were studied. P. chrysosporium could grow in liquid media with400mg L-1Pb(II), which might be due to its two possible responses:dense and tight twist of hyphae, and secretion from mycelia to resist Pb. During solid-state fermentation, fungal colonization capability was positively correlated with the removal efficiency of soluble-exchangeable Pb when its content was higher than8.2mg kg"1dry mass. Carboxymethyl cellulase activity and cellulose degradation were inhibited at different initial Pb concentrations, whereas low initial Pb concentrations strengthened xylanase and ligninolytic-enzyme activities and the hemicellulose and lignin degradation.(2) Degradation of Pb-polluted lignocellulosic waste and the restrain of Pb hazards by P. chrysosporium were studied. P. chrysosporium degraded43.1%of lignocellulose, formed humus and reduced active Pb ions, even at the concentration of400mg kg-1dry mass of Pb. The highest lignocellulose degradation (56.8%) and organic matter loss (64.0%) were found at Pb concentration of30mg kg-1, and selective lignin biodegradation was enhanced. Scanning electron micrographs with energy spectra showed that Pb was immobilized via two possible routes:adsorption and cation exchange on hypha, and the chelation by fungal metabolite.(3) Treatment of Pb-polluted solid waste by composting with ligninolytic fungi was studied. Microbial biomass and CO2production increased, whereas soluble-exchangeable Pb content was reduced to0%. water-soluble organic C/N ratio might be not a suitable evaluating indicator for the composting of metal-polluted waste, while lignin and coarse fibre were suggested.(4) Pb-polluted soil could be remediated by composting with ligninolytic fungi, and microbial activities and carbon source utilization were enhanced. Active Pb form was transformed into the residual and Fe-Mn oxidation forms. It might be due to adsorption on fungi mycelia and complexation of humus.
The four section focuses on the model construction for relationship between enzyme activities, microbial community and lignin degradation respectively.(2) It was confirmed that ANN model could be used to predicted lignin degradation by the input of LiP and MnP activities.(2) ANN model could predicted lignin degradation by the input of community information. These showed the advantage of ANN application in complex relationship resolution, which could be a reference for quantification of effect of enzymes and microorganisms on lignin degradation.
This dissertation revealed microbial community succession and the mechanisms of solid waste composting by ligninolytic microorganisms, which could provide microbial information for the understanding of composting process and benefit the studies of complex microbial inoculants. It is expected to lay the foundation for the development of a new composting method with high efficiency, which could conquer some deficiency existed in traditional composting technology.
本文探讨了堆肥过程中微生物群落的演替规律,系统地研究了高效木质素降解功能微生物与其胞外酶的行为机制及对堆肥微生物群落结构与活性、木质素降解、腐殖质形成的作用机理,并考察强化堆肥法对堆肥重金属毒性的影响,发展了新型高效的基于木质素降解功能微生物、复合酶的强化堆肥技术与理论。本文的具体研究工作及成果包括以下4个部分内容：
第1部分为堆肥过程微生物群落结构变化与木质纤维素降解的相关性研究。(1)采用醌指纹法研究了堆肥过程中微生物群落的演替。发现微生物群落结构发生了明显改变。高温阶段,Q-9(H2)、Q-10(H2)表征的真菌是半纤维素、纤维素的主要降解者,MK-7、Q-10分别表征的细菌、嗜热真菌是主要木质素降解者,而Q-9表征的黑曲霉、青霉等嗜温木质素降解真菌量少,降解作用有限。降温阶段,Q-9和长链甲基萘醌增多,其与优势醌MK-7分别表征的真菌、放线菌、细菌协同作用,促使木质素大量降解,并呈较高的纤维素降解率。(2)含MK-9(H2)的放线菌分别与MK-9、MK-9(H6)、MK-10(H6)表征的放线菌有良好共存性,含MK-5(H2)的细菌分别与MK-9、MK-9(H2)表征的放线菌友好共存,互促生长繁殖。限制性片段长度多态性分析显示,细菌在高温期最活跃,堆肥过程细菌群落基因多样性变化小。(3)结合主成分、相关性分析等探讨了木质素降解微生物群落的组成及其与木质纤维素降解的相关性。发现Q-9、MK-7、MK-8(H2)、MK-9、 MK-9(H2)、MK-9(H6)、MK-9(H8)和MK-10(H6)表征的嗜温真菌、细菌和放线菌组成的特定群落是起关键作用的木质素降解群落,与木质素降解正相关,含后7种醌的微生物群落还与纤维素降解正相关。(4)证实了木质素的降解显著影响半纤维素、纤维素降解,指出堆肥过程木质素降解真菌的劣势地位限制了木质素的降解,可通过接种木质素降解真菌促进木质素降解、提高堆肥效率。
第2部分着重研究木质素降解功能微生物、胞外酶强化堆肥技术及机理。(1)针对高效木质素降解功能微生物白腐真菌特性与降解机理展开研究。发现温度和含水率是影响该菌固态发酵产酶和降解能力的最主要因素,确定最佳工艺参数：菌种驯化5d,接种量0.8%,吐温80添加量0.3%,温度37℃,含水率85%。证实该菌能严重破坏木质素结构,使苯环开环,难降解的大分子长键烃断裂成易降解的小分子短键烃。提出了用关联分析思想判别白腐菌产酶过程最显著影响因素,建立了GM(1,1)模型预测日产酶量,发现在最显著影响因素含水率不变时,GM(1,1)模型能快速准确地预测产酶,减少实验工作量。(2)建立了木质素降解功能微生物强化堆肥法处理木质纤维素类废物。证实强化堆肥法可增加微生物活性,提高木质素降解率至43.9%,使有机物降解更彻底。(3)进一步研究强化堆肥过程中的微生物群落演替和腐殖质形成。发现强化堆肥法显著提高了成品腐殖化程度(54.1%)和总微生物量,但降低了群落多样性指数、均匀性指数,使Q-9、长链甲基萘醌分别表征的木质素降解真菌、放线菌增多,尤其木质素降解真菌成为优势菌,Q-8、MK-7分别表征的固氮细菌、木质素降解细菌增多,而其它微生物减少。聚类分析和主成分分析显示,强化堆肥法明显改变了微生物群落结构组成,尤其在16～28d影响最大,大大加强了木质素降解真菌、放线菌在整个堆肥体系中的影响力,利于木质素降解。(4)制备了木质素降解功能复合酶,考察了其在稻草、米糠、蔬菜残余、土壤4种典型堆肥基质表面的传输行为与附着性能,并研究其降解天然木质素机理。发现复合酶在稻草、米糠中传输性能最佳,可传输至基质柱深层；在4种基质表面的附着都属有利吸附,利于酶与基质的有效接触。复合酶酶解木质素时,通过氧化苯甲醇单元、脱甲氧基和甲基、断裂部分碳碳双键、醚键与Ca-Cβ键并氧化Ca侧链成羧酸、芳环开环及芳环取代等一系列反应,导致木质素结构中醇羟基、亚甲基、甲氧基、甲基、醚键、碳碳双键、苯环等减少,香草酸、阿魏酸等酯型结构被分解,木质素二聚体解聚成低分子物质,羰基、取代芳环等增多。(6)研究了功能复合酶强化堆肥法对微生物群落碳源代谢、木质素降解的影响。结果显示,添加复合酶显著增强了总有机质的降解和微生物群落碳源代谢能力。Biolog分析表明,复合酶主要影响了微生物群落对17种碳源的利用,大大提高了丙酮酸甲酯、a-环式糊精、D-甘露醇、D-半乳糖醛酸、衣康酸、L-天冬酰胺的利用,降低了D,L-a-磷酸甘油、L-苏氨酸、甘氨酰-L-谷氨酸、腐胺的利用。复合酶显著促进了木质素降解,可能与其既能催化木质素分解,又能增强微生物对酚类化合物、羧酸类等木质素降解中间产物的代谢有关。
第3部分为木质素降解功能微生物强化堆肥法处理Pb污染基质及过程机理研究。(1)探讨了Pb胁迫下木质素降解功能微生物白腐菌的反应行为和降解能力。结果表明,该菌通过菌丝缠绕更致密及分泌胞外物质2种反应行为来抵御Pb毒性,在含400mg L-1Pb2+的液态培养基中能较好生长。Pb污染固态基质处理中,白腐菌定殖能力与水溶交换态Pb减少率(含量高于8.2mg kg-1时)正相关；不同浓度Pb对纤维素酶活、纤维素降解有抑制作用,而低浓度Pb使木聚糖酶、木质素降解酶活性和半纤维素、木质素降解增强。(2)系统研究了黄孢原毛平革菌处理不同浓度Pb污染木质纤维素废物的整个过程,探讨了它对Pb的钝化机制。发现即使初始Pb浓度为400mg kg-1干基质时,该菌仍能降解43.1%的木质纤维素,并形成腐殖质,大大减少游离态Pb。初始Pb浓度为30mg kg-1时,木质纤维素、有机质降解率高达56.8%、64.0%,木质素降解选择性指数较高。经扫描电镜与能谱分析发现,该菌对Pb的钝化可能主要通过：菌丝表面的吸附与离子交换作用,及该菌代谢物的络合作用。(3)研究了木质素降解功能微生物强化堆肥法处理Pb污染垃圾及机理。发现强化堆肥法能经济有效地处理Pb污染垃圾,堆制过程中有较高的微生物量、CO2释放量,使成品中水溶交换态Pb含量为0%、毒性小。且指出水溶性有机C/N变化不宜作为重金属垃圾堆肥的腐熟度评价指标,可考虑改用木质素、粗纤维余量。(4)研究了木质素降解功能微生物强化堆肥法修复Pb污染土壤及机理。发现由于微生物菌丝体对Pb离子的强吸附作用及堆肥过程中生成腐殖质的络合作用,污染土壤经强化堆肥法修复后,土壤微生物活性和群落碳源代谢能力增强,游离态Pb转为以残留态和铁锰氧化物结合态存在。
第4部分着重开展关键生物酶活性及微生物群落演替与木质素降解的映射模型的构建。(1)构建了木质纤维素废物处理体系中酶活与木质素降解率关系的人工神经网络(ANN)模型,该模型中输入酶活值能准确预测木质素的降解。(2)构建了堆肥过程中微生物群落演替与木质素降解率关系的ANN模型,该模型中输入群落信息能有效预测木质素降解。ANN模型在复杂关系解析中具优越性,为定量分析关键酶、微生物群落对木质素降解的影响提供了技术支持,可高效指导有关复合酶、复合菌剂的开发和应用。
本论文研究揭示了堆肥微生物群落演替及木质素降解功能微生物强化堆肥机理,可为科学认识堆肥反应本质提供微生物群落信息,为微生物选育、复合生物促进剂等相关研究的合理开展提供依据；并为高效快速堆肥新方法的发展奠定基础,以克服传统堆肥法处理效率低、腐熟程度差等不足。
Composting, as an economic and effective technology for solid waste treatment, is one of important research subjects. But the application of traditional composting are limited due to the biodegradative recalcitrance of lignin and the stresss of heavy metal in composting materials. So the improvement of composting efficiency and heavy metal control are very important. Since microbial community are responsible for organic matter degradation during composting, the study on microbial community succession is a basis of composting technology and theory researches. To improve composting efficiency and control heavy metal pollution, exploring a new composting technology and theory based on microbial community study are needed. It would promote the development of composting technology, and be of theoretical and practical value for efficient and rapid composting treatment.
This dissertation discussed microbial community succession, and investigated the effects of ligninolytic microorganisms and enzymes on microbial activities, lignin degradation and humus formation in composting. The control of heavy metal are also considered in this study. According to these researches, we developed a new and efficient composting techonology based on ligninolytic microorganisms or enzymes. This dissertation is composed of four sections.
The first section describes the research on the relationship between microbial community successtion and lignocellulose degradation.(1) Microbial populations and their relationship to bioconversion during composting were studied by quinone profiling. The obvious changes in microbial population were observed. Fungi indicated by Q-9(H2) and Q-10(H2) were considered to be the most important hemicellulose and cellulose-degrading microorganisms during thermophilic stage. MK-7and Q-10, indicative of certain bacteria and fungi respectively, were responsible for lignin degradation at the thermophilic stage, whereas the mesophile ligninolytic fungi (Aspergillus niger, Penicillium, and etc.) indicated by Q-9is limited. The highest lignin degradation ratio and good cellulose degradation were found at the cooling stage and were correlated with Q-9, MK-7and long-chain menaquinones attributed to mesophilic fungi, bacteria and actinomycetes, respectively.(2) MK-9(H2) positively correlated with MK-9MK-9(H6) and MK-10(H6) respectively, which suggests coexistence and possibly even cooperativity among the corresponding microorganisms. Significant positive correlation was also suggested for the bacteria indicated by MK-5(H2) and the actinomycetes containing MK-9or MK-9(H2) as major quinone. RFLP analysis showed there were no obvious changes in genetic diversity of bacterial community during composting, and bacterial community is very active during thermophilic stage.(3) Principal component and correlation analyses showed a significant positive correlation between lignin degradation ratio and the total content of Q-9、MK-7、MK-8(H2)、MK-9、MK-9(H2)、MK-9(H6)、MK-9(H8)和MK-10(H6), which indicated that the microbial community consisting of mesophilic fungi, bacteria and actinomycetes played a key role in lignin degradation. And a positive correlation between the latter7quinones and cellulose degradation was found.(4) Lignin degradation significantly affected hemicellulose and cellulose degradation. The inferior position of ligninolytic fungi in composting might be responsible for slow degradation of lignin. It is expected that inoculating ligninolytic fungi would accelerate lignin degradation and promote composting efficiency.
The second section focuses on the study of mechanisms of the composting technology by adding ligninolytic microorganism or enzymes.(1) The characteristics and degradation ability of ligninolytic fungus (white-rot fungus) were studied. Temperature and substrate moisture were confirmed to be the main effect factors. The optimal treament conditions are:5days of fungi incubation,0.8%of inoculum,0.3%of Tween80addition,37℃and85%of water content of substrate. This fungus could destroy lignin structure by the cleavage of aromatic ring and the transformation of long-bond hydrocarbons of giant molecule to the short-bond. The study also applied relational analysis to the enzyme production process of white-rot fungi, and constructed GM(1,1) model which could be used to predicted enzyme production to reduce experimental quantity.(2) Composting of lignocellulosic waste by ligninolytic fungi was studied. By this composting method, microbial activities were enhanced, and lignin degradation ratio reached43.9%, and more completed degradation of organic matter was also found.(3) The higher humification degree (54.1%) and total microbial biomass were observed during the composting process with the inocula of ligninolytic fungi than those in tradition composting. While the low diversity and evenness of microbial community were found. It might be because the lignin-degrading fungi and actinomycetes respectively indicated by Q-9and long-chain menaquinones increased, and bacteria with nitrogen fixation and lignin-degrading bacteria indicated by Q-8and MK-7increased, whereas other microorganisms decreased. Composting method with fungi inocula changed microbial community structure obviously, especially from day16to28, enhanced the effect of lignin-degrading fungi and actinomycetes.(4) The transfer and adsorption behaviours and degradation mechanisms of ligninolytic enzymes (LiP-MnP) were studied. The transfer abilities of ligninolytic enzymes on straw and chaff are better than vegetable residues and soil. The adsorption on these four substrates belonged to favourable adsorption, which ensured the effective contact between enzymes and substrates. Degradation mechanisms of ligninolytic enzymes included:benzyl alcohol unit oxidation, methoxy and methyl group removal, C=C and ether bond and Ca-Cβ cleavage, and aromatic ring cleavage and replacement. Therefore, alcohol hydroxyl,-CH2,-OCH3,-CH3, ether bond, C=C, and aromatic ring in lignin decreased. Vanillic acid and ferulic acid unit in lignin were decomposed, and lignin dimers were degraded into low-weight matters.(6) Carbon source utilization and lignin degradation by microbial community were studied during composting with LiP-MnP enzymes. Addition of LiP-MnP enzymes significantly promoted organic matter degradation and carbon source utilization. It was found by Biolog analysis that the enzymes mainly affected the utilization of17carbon source. The utilization of pyruvic acid methyl ester, a-cyclodextrin, D-mannitol, D-galacturonic acid, itaconic acid and L-asparagine were enhanced, whereas the utilization of D,L-a-glycerol phosphate, L-threonine, glycyl-L-glutamic acid and putrescine were weakened. Lignin degradation was significantly promoted by LiP-MnP enzymes. There might be two reasons:the enzymes catalyzed lignin degradation, and enhanced the utilization of lignin degradation products such as phenolic compound and carboxylic acid.
The third section describes the research on composting of Pb-polluted substrates by inocula of ligninolytic fungi.(1) Mycelial growth and the degradation of lignocellulosic waste by ligninolytic fungus Phanerochaete chrysosporium under lead stress were studied. P. chrysosporium could grow in liquid media with400mg L-1Pb(II), which might be due to its two possible responses:dense and tight twist of hyphae, and secretion from mycelia to resist Pb. During solid-state fermentation, fungal colonization capability was positively correlated with the removal efficiency of soluble-exchangeable Pb when its content was higher than8.2mg kg"1dry mass. Carboxymethyl cellulase activity and cellulose degradation were inhibited at different initial Pb concentrations, whereas low initial Pb concentrations strengthened xylanase and ligninolytic-enzyme activities and the hemicellulose and lignin degradation.(2) Degradation of Pb-polluted lignocellulosic waste and the restrain of Pb hazards by P. chrysosporium were studied. P. chrysosporium degraded43.1%of lignocellulose, formed humus and reduced active Pb ions, even at the concentration of400mg kg-1dry mass of Pb. The highest lignocellulose degradation (56.8%) and organic matter loss (64.0%) were found at Pb concentration of30mg kg-1, and selective lignin biodegradation was enhanced. Scanning electron micrographs with energy spectra showed that Pb was immobilized via two possible routes:adsorption and cation exchange on hypha, and the chelation by fungal metabolite.(3) Treatment of Pb-polluted solid waste by composting with ligninolytic fungi was studied. Microbial biomass and CO2production increased, whereas soluble-exchangeable Pb content was reduced to0%. water-soluble organic C/N ratio might be not a suitable evaluating indicator for the composting of metal-polluted waste, while lignin and coarse fibre were suggested.(4) Pb-polluted soil could be remediated by composting with ligninolytic fungi, and microbial activities and carbon source utilization were enhanced. Active Pb form was transformed into the residual and Fe-Mn oxidation forms. It might be due to adsorption on fungi mycelia and complexation of humus.
The four section focuses on the model construction for relationship between enzyme activities, microbial community and lignin degradation respectively.(2) It was confirmed that ANN model could be used to predicted lignin degradation by the input of LiP and MnP activities.(2) ANN model could predicted lignin degradation by the input of community information. These showed the advantage of ANN application in complex relationship resolution, which could be a reference for quantification of effect of enzymes and microorganisms on lignin degradation.
This dissertation revealed microbial community succession and the mechanisms of solid waste composting by ligninolytic microorganisms, which could provide microbial information for the understanding of composting process and benefit the studies of complex microbial inoculants. It is expected to lay the foundation for the development of a new composting method with high efficiency, which could conquer some deficiency existed in traditional composting technology.
引文
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