农业废物堆肥化中木质素的降解及其微生物特性研究
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摘要
木质素是农业废物堆肥化过程中的主要限速有机物,其降解被认为是快速堆肥的关键。本文系统研究了农业废物堆肥化中木质素的降解过程及其微生物特性,首先分别运用传统方法、Biolog法、PLFA法、醌指纹法研究木质素降解微生物群落演替规律,其次筛选高效优势的木质素降解微生物,并研究非白腐真菌简青霉(P.simplicissimum)、黑曲霉(A.niger)的木质素降解特性和降解机理,最后研究一株高效优势的木质素降解放线菌栗褐链霉菌对腐殖质产生的作用。
传统方法研究发现,木质素降解主要发生在高温期和二次发酵期。多酚氧化酶PPO和过氧化物酶POD是木质素降解的主要酶系,其中PPO的催化氧化作用较强。木质素降解细菌、放线菌、真菌在堆肥化各阶段变化复杂,其中,放线菌是主要的木质素降解者,从整个过程看,各类群微生物对木质素降解的贡献为放线菌>真菌>细菌。
细菌群落在堆肥化进程中木质素的生物降解方面起着重要的协作作用。应用Biolog法检测堆肥化过程中细菌群落沿替,并通过多元统计分析,发现堆肥初期细菌生长快速、群落丰富,随着堆制的进行,其平均活性逐渐下降。细菌群落结构在一次发酵期间发生着剧烈变化,二次发酵期间趋于稳定。能转化BiologGN2板上第一、二、四类碳源的细菌与木质纤维素的转化有关,其中第一、二类碳源表征农业废物堆肥化进程中的主要细菌种群,第四类碳源表征细菌能够耐受高温。
应用PLFA-PLS法构建了木质素含量与PLFA之间的定量回归模型,分析模型参数可发现,农业废物堆肥化过程中木质素的有效降解是数量少、能力强与数量多、能力弱的微生物共同作用的结果,前者主要是真菌、放线菌,后者主要是细菌,其中真菌与放线菌在堆肥化过程中占主导地位。微生物种类较之数量对于木质素降解更为有利。在高温期,有效的木质素降解微生物群落组成为:革兰氏阳性细菌/革兰氏阴性细菌/放线菌/真菌=42/35/6/17;在二次发酵期为:革兰氏阳性细菌/革兰氏阴性细菌/放线菌/真菌=58/23/4/15。微生物浓度数量级为108cells/gdw。醌指纹法证实堆肥化过程中发生着剧烈的群落演替,且微生物群落
结构随着堆肥成熟逐步趋于复杂。MK-7指示的细菌和Q-9指示的真菌在堆肥化初期为优势菌群,主要降解易降解有机物,与木质纤维素降解无关。部分饱和、长链的甲基萘醌指示的放线菌是高温期优势微生物,这类微生物是木质素降解的主要作用者。放线菌、真菌和小部分细菌在二次发酵期间共同协作,对木质素降解有一定作用。Q-9与Q-10(H2)指示的微生物是纤维素与半纤维素降解的主要微生物,而木质素的有效降解是一些还未被广泛认识的含Q-9(H2)为主要醌的微生物与高温放线菌共同协作的结果。
筛选到几株较强的木质素降解菌,包括AspergillusnigerF4-1,AspergillusoryzaeF1-4、PenicilliumsimplicissimumF1-1、StreptomycesbadiusA4-2以及高温放线菌Thermoactinomycessp.A2-2。对黑曲霉的研究发现该菌能攻击木质素酚型、非酚型结构和苯环侧链结构。在实验条件下,该菌主要依靠胞外LiP、Lac、MnP共同作用降解木质素,并表现出次级代谢降解机制。简青霉具有较强的攻击木质素酚型结构的能力,降解木质素主要依靠胞外Lac,表现出明显的初级代谢机制。pH值对降解系统有较大的影响,Mn2+不利于降解,而Cu2+对木质素降解有较强的促进作用。
对简青霉产酶调控机理研究发现,附加碳源抑制漆酶Lac活性,适量氮源可显著提高酶活,常用的Lac诱导剂ABTS和二甲苯胺并不能促进简青霉产生Lac。在发酵过程中,前期Lac主要起聚合作用,而后期可能在某些降解产物的协同作用下,Lac表现出较好的降解能力。各种附加营养条件诱使简青霉在第3d出现Lac活性峰,该峰值下漆酶聚合能力特别强,直接影响最后木质素的降解率。
比较黄孢原毛平革菌和栗褐链霉菌对腐殖质形成的作用,结果表明,接种木质素降解菌能促进发酵体系中腐殖质的形成,栗褐链霉菌的促进作用更强。由于黄孢原毛平革菌和栗褐链霉菌具有不同的木质素降解机制,它们分别形成腐殖质的途径也不同。黄孢原毛平革菌转化木质素生成简单分子(FA),然后聚合FA生成复杂的分子(HA),而栗褐链霉菌改性木质素,先生成HA,然后将HA转化成FA。
Since lignin is the major organic compound that limiting composting speed, its degradation is essential for the operation of composting. Lignin biodegradation and its microbial characteristics during composting of agricultural wastes were studied. Firstly, traditional method, Biolog assay, PLFA analysis and quinone profiles were used respectively to investigate the microbial community succession with lignin degrading potential. Secondly, dominant and effectively lignin degrading microorganisms were screened. Nonwhite-rot fungi, Penicillium simplicissimum and Aspergillus niger, were found to be important in lignin degradation. Their degrading characteristics and mechanisms were also studied. Finally, effect of lignin biodegradation on humification by Streptomyces badius isolated from maturation phase was investigated.
The results obtained through the traditional method indicated that lignin was mainly degraded during the thermophilic phase and maturation phase of composting. Polyphenol oxidase (PPO) and peroxidase (POD), especially PPO was found to be the most important catalyzer in lignin degradation. Bacteria, fungi and actinomycetes with lignin degrading ability changed greatly during every compost phase. Actinomycetes were found to be the most effective microorganism to degrade lignin followed by fungi and bacteria.
Since bacterial communities played important synergic function in lignin degradation, they were analyzed using Biolog method in agricultural waste composting. The results of cluster analysis and principle component analysis indicated that bacterial communities varied greatly during the first stage of composting, while began to stabilize during the second stage. Bacteria that could utilize the first, second and fourth kinds of carbon sources on Biolog GN2 plate were found to be able to degrade lignocellulose. Thereinto, the first and second kinds of carbon sources were the dominant ones during composting, while the bacteria that metabolize the fourth kind of carbon sources were believed to be thermophilic ones.
Phospholipid fatty acid (PLFA) and Partial Least- squares Regression (PLS) methods were selected to establish the quantitative regressive model
between lignin content and PLFA. Through the analysis of model parameters, it was found that a few microorganisms with strong lignin degrading ability and plentiful microbes with weak lignin degrading ability cooperated during the composting of agricultural wastes. The former were fungi and actinomycetes that dominant in lignin degradation, while the latter was bacteria. Microbial species were believed to be more important than amounts. During the thermophilic phase, the effective lignin degrading microbial proportion was 42:35:6:17 (Gram-positive bacteria: Gram-negative bacteria: actinomycetes: fungi), while the proportion was 58:23:4:15 during the maturation phase of composting. The microbial content was about 108cells/g dw.
The changes of microbial community during agricultural waste composting were successfully studied by quinone profiles. Mesophilic bacteria indicated by MK-7 and mesophilic fungi containing Q-9 as major quinone were predominant and seemed to be important during the initial stage of composting. Actinobacteria indicated by a series of partially saturated and long-chain menaquinones were preponderant during the thermophilic period. While actinomycetes, fungi and some bacteria, especially those microbes containing MK-7(H4) found in Gram-positive bacteria with a low G+C content or actinomycetes were found cooperate during the latter maturating period. Since lignocellulsoe is abundant in the agricultural wastes and its degradation is essential for the operation of composting, it’s important to establish the correlation between the quinone profiles changes and lignocellulose degradation. The microbes containing Q-9 or Q-10(H2) as major quinone were found to be the most important hemicellulose and cellulose degrading microorganisms during composting. While the microorganisms containing Q-9(H2) as major quinone and many thermophilic actinomycetes were believed to be responsible for lignin degradation during agricultural waste composting.
Several strains include Aspergillus niger F4-1, Aspergillus oryzae F1-4, Penicillium simplicissimum F1-1, Streptomyces badiusA4-2 and thermophilic actinomycetes Thermoactinomyces sp.A2-2 that have strong lignin degrading ability were obtained through the screening experiments. A. niger was capable of utilizing phenolic and nonphenolic lignin model compounds. Lignin peroxidase (LiP), Manganses peroxidase (MnP), Laccase were believed to be the most important catalyzes in biodegrading process. P. simplicissimum was capable of utilizing several lignin model compounds, making aromatic dyes decolourise and degrading natural lignin. All these results proved that P. simplicissimum has ligninolytic ability. Laccase was the most important catalyzes in the biodegrading process. Different from the degrading mechanism of the white-rot fungi, the lignin degradation by P.simplicissimum mainly happened during the primary metabolism and it was greatly influenced by the pH of media, the concentration of Cu2+ and Mn2+.
The laccase activities of P. simplicissimum during solid state fermentation with rice straw were studied. And the degradation of lignocellulose was also determined. Results show that all supplemental carbon sources inhibited the laccase activity in different degrees. While proper concentration of supplemental nitrogen sources remarkably enhanced the laccase activity. The enhancement of laccase activity by ordinary laccase inducers 2, 2’-azino-bis (3-ethylbenzthiazoline- 6-sulfonic acid) and xylidine was not observed in this study. Lignocellulose degradation was improved when laccase activity was relatively low during prophase of fermentation, which proved the polymerizing function of laccase in lignin degradation by P. simplicissimum. Some metabolites may act as mediators to help lignin degradation during the fermentation anaphase. All supplemental sources may induce the peak of Lac activity on day 3 when the polymerizing function was especially strong.
The effect of lignin biodegradation on humification by Phanerochaete chrysosporium and S. badius were compared. The results indicate that lignin degrading microorganisms could promote humus formation. Since the degrading mechanisms of two strains were different, the approaches of humus formation from lignin with P. chrysosporium and S. badius were also different. P. chrysosporium metabolize lignin to simple molecules (FA), which then were polymerized to complex molecules (HA). While S. badius modified lignin to HA, which then were transformed to FA.
传统方法研究发现,木质素降解主要发生在高温期和二次发酵期。多酚氧化酶PPO和过氧化物酶POD是木质素降解的主要酶系,其中PPO的催化氧化作用较强。木质素降解细菌、放线菌、真菌在堆肥化各阶段变化复杂,其中,放线菌是主要的木质素降解者,从整个过程看,各类群微生物对木质素降解的贡献为放线菌>真菌>细菌。
细菌群落在堆肥化进程中木质素的生物降解方面起着重要的协作作用。应用Biolog法检测堆肥化过程中细菌群落沿替,并通过多元统计分析,发现堆肥初期细菌生长快速、群落丰富,随着堆制的进行,其平均活性逐渐下降。细菌群落结构在一次发酵期间发生着剧烈变化,二次发酵期间趋于稳定。能转化BiologGN2板上第一、二、四类碳源的细菌与木质纤维素的转化有关,其中第一、二类碳源表征农业废物堆肥化进程中的主要细菌种群,第四类碳源表征细菌能够耐受高温。
应用PLFA-PLS法构建了木质素含量与PLFA之间的定量回归模型,分析模型参数可发现,农业废物堆肥化过程中木质素的有效降解是数量少、能力强与数量多、能力弱的微生物共同作用的结果,前者主要是真菌、放线菌,后者主要是细菌,其中真菌与放线菌在堆肥化过程中占主导地位。微生物种类较之数量对于木质素降解更为有利。在高温期,有效的木质素降解微生物群落组成为:革兰氏阳性细菌/革兰氏阴性细菌/放线菌/真菌=42/35/6/17;在二次发酵期为:革兰氏阳性细菌/革兰氏阴性细菌/放线菌/真菌=58/23/4/15。微生物浓度数量级为108cells/gdw。醌指纹法证实堆肥化过程中发生着剧烈的群落演替,且微生物群落
结构随着堆肥成熟逐步趋于复杂。MK-7指示的细菌和Q-9指示的真菌在堆肥化初期为优势菌群,主要降解易降解有机物,与木质纤维素降解无关。部分饱和、长链的甲基萘醌指示的放线菌是高温期优势微生物,这类微生物是木质素降解的主要作用者。放线菌、真菌和小部分细菌在二次发酵期间共同协作,对木质素降解有一定作用。Q-9与Q-10(H2)指示的微生物是纤维素与半纤维素降解的主要微生物,而木质素的有效降解是一些还未被广泛认识的含Q-9(H2)为主要醌的微生物与高温放线菌共同协作的结果。
筛选到几株较强的木质素降解菌,包括AspergillusnigerF4-1,AspergillusoryzaeF1-4、PenicilliumsimplicissimumF1-1、StreptomycesbadiusA4-2以及高温放线菌Thermoactinomycessp.A2-2。对黑曲霉的研究发现该菌能攻击木质素酚型、非酚型结构和苯环侧链结构。在实验条件下,该菌主要依靠胞外LiP、Lac、MnP共同作用降解木质素,并表现出次级代谢降解机制。简青霉具有较强的攻击木质素酚型结构的能力,降解木质素主要依靠胞外Lac,表现出明显的初级代谢机制。pH值对降解系统有较大的影响,Mn2+不利于降解,而Cu2+对木质素降解有较强的促进作用。
对简青霉产酶调控机理研究发现,附加碳源抑制漆酶Lac活性,适量氮源可显著提高酶活,常用的Lac诱导剂ABTS和二甲苯胺并不能促进简青霉产生Lac。在发酵过程中,前期Lac主要起聚合作用,而后期可能在某些降解产物的协同作用下,Lac表现出较好的降解能力。各种附加营养条件诱使简青霉在第3d出现Lac活性峰,该峰值下漆酶聚合能力特别强,直接影响最后木质素的降解率。
比较黄孢原毛平革菌和栗褐链霉菌对腐殖质形成的作用,结果表明,接种木质素降解菌能促进发酵体系中腐殖质的形成,栗褐链霉菌的促进作用更强。由于黄孢原毛平革菌和栗褐链霉菌具有不同的木质素降解机制,它们分别形成腐殖质的途径也不同。黄孢原毛平革菌转化木质素生成简单分子(FA),然后聚合FA生成复杂的分子(HA),而栗褐链霉菌改性木质素,先生成HA,然后将HA转化成FA。
Since lignin is the major organic compound that limiting composting speed, its degradation is essential for the operation of composting. Lignin biodegradation and its microbial characteristics during composting of agricultural wastes were studied. Firstly, traditional method, Biolog assay, PLFA analysis and quinone profiles were used respectively to investigate the microbial community succession with lignin degrading potential. Secondly, dominant and effectively lignin degrading microorganisms were screened. Nonwhite-rot fungi, Penicillium simplicissimum and Aspergillus niger, were found to be important in lignin degradation. Their degrading characteristics and mechanisms were also studied. Finally, effect of lignin biodegradation on humification by Streptomyces badius isolated from maturation phase was investigated.
The results obtained through the traditional method indicated that lignin was mainly degraded during the thermophilic phase and maturation phase of composting. Polyphenol oxidase (PPO) and peroxidase (POD), especially PPO was found to be the most important catalyzer in lignin degradation. Bacteria, fungi and actinomycetes with lignin degrading ability changed greatly during every compost phase. Actinomycetes were found to be the most effective microorganism to degrade lignin followed by fungi and bacteria.
Since bacterial communities played important synergic function in lignin degradation, they were analyzed using Biolog method in agricultural waste composting. The results of cluster analysis and principle component analysis indicated that bacterial communities varied greatly during the first stage of composting, while began to stabilize during the second stage. Bacteria that could utilize the first, second and fourth kinds of carbon sources on Biolog GN2 plate were found to be able to degrade lignocellulose. Thereinto, the first and second kinds of carbon sources were the dominant ones during composting, while the bacteria that metabolize the fourth kind of carbon sources were believed to be thermophilic ones.
Phospholipid fatty acid (PLFA) and Partial Least- squares Regression (PLS) methods were selected to establish the quantitative regressive model
between lignin content and PLFA. Through the analysis of model parameters, it was found that a few microorganisms with strong lignin degrading ability and plentiful microbes with weak lignin degrading ability cooperated during the composting of agricultural wastes. The former were fungi and actinomycetes that dominant in lignin degradation, while the latter was bacteria. Microbial species were believed to be more important than amounts. During the thermophilic phase, the effective lignin degrading microbial proportion was 42:35:6:17 (Gram-positive bacteria: Gram-negative bacteria: actinomycetes: fungi), while the proportion was 58:23:4:15 during the maturation phase of composting. The microbial content was about 108cells/g dw.
The changes of microbial community during agricultural waste composting were successfully studied by quinone profiles. Mesophilic bacteria indicated by MK-7 and mesophilic fungi containing Q-9 as major quinone were predominant and seemed to be important during the initial stage of composting. Actinobacteria indicated by a series of partially saturated and long-chain menaquinones were preponderant during the thermophilic period. While actinomycetes, fungi and some bacteria, especially those microbes containing MK-7(H4) found in Gram-positive bacteria with a low G+C content or actinomycetes were found cooperate during the latter maturating period. Since lignocellulsoe is abundant in the agricultural wastes and its degradation is essential for the operation of composting, it’s important to establish the correlation between the quinone profiles changes and lignocellulose degradation. The microbes containing Q-9 or Q-10(H2) as major quinone were found to be the most important hemicellulose and cellulose degrading microorganisms during composting. While the microorganisms containing Q-9(H2) as major quinone and many thermophilic actinomycetes were believed to be responsible for lignin degradation during agricultural waste composting.
Several strains include Aspergillus niger F4-1, Aspergillus oryzae F1-4, Penicillium simplicissimum F1-1, Streptomyces badiusA4-2 and thermophilic actinomycetes Thermoactinomyces sp.A2-2 that have strong lignin degrading ability were obtained through the screening experiments. A. niger was capable of utilizing phenolic and nonphenolic lignin model compounds. Lignin peroxidase (LiP), Manganses peroxidase (MnP), Laccase were believed to be the most important catalyzes in biodegrading process. P. simplicissimum was capable of utilizing several lignin model compounds, making aromatic dyes decolourise and degrading natural lignin. All these results proved that P. simplicissimum has ligninolytic ability. Laccase was the most important catalyzes in the biodegrading process. Different from the degrading mechanism of the white-rot fungi, the lignin degradation by P.simplicissimum mainly happened during the primary metabolism and it was greatly influenced by the pH of media, the concentration of Cu2+ and Mn2+.
The laccase activities of P. simplicissimum during solid state fermentation with rice straw were studied. And the degradation of lignocellulose was also determined. Results show that all supplemental carbon sources inhibited the laccase activity in different degrees. While proper concentration of supplemental nitrogen sources remarkably enhanced the laccase activity. The enhancement of laccase activity by ordinary laccase inducers 2, 2’-azino-bis (3-ethylbenzthiazoline- 6-sulfonic acid) and xylidine was not observed in this study. Lignocellulose degradation was improved when laccase activity was relatively low during prophase of fermentation, which proved the polymerizing function of laccase in lignin degradation by P. simplicissimum. Some metabolites may act as mediators to help lignin degradation during the fermentation anaphase. All supplemental sources may induce the peak of Lac activity on day 3 when the polymerizing function was especially strong.
The effect of lignin biodegradation on humification by Phanerochaete chrysosporium and S. badius were compared. The results indicate that lignin degrading microorganisms could promote humus formation. Since the degrading mechanisms of two strains were different, the approaches of humus formation from lignin with P. chrysosporium and S. badius were also different. P. chrysosporium metabolize lignin to simple molecules (FA), which then were polymerized to complex molecules (HA). While S. badius modified lignin to HA, which then were transformed to FA.
引文
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