原生质体技术筛选刺芹侧耳高产多糖和漆酶菌株的研究
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摘要
真菌多糖和漆酶作为真菌的两大类重要生理活性物质,越来越受到人们的关注。真菌多糖作为“生物效应调节剂”,具有很强的非特异性免疫增强作用,通过多方面机制实现抗肿瘤活性。目前已有众多真菌多糖在临床上得以应用,用来治疗免疫缺陷性疾病、肿瘤、病毒感染等相关疾病。此外,真菌多糖还具有抗氧化、降血糖、降血脂等多种药理活性,已被开发为功能性食品添加剂使用。真菌漆酶是一种重要的木质素降解酶,其作用的底物之多,使其在环保、食品、造纸、纺织等工业均具有良好的应用。
刺芹侧耳(Pleurotus eryngii),是20世纪90年代我国引种成功的一种珍稀食药用真菌,因其据有杏仁的香味,菌肉肥厚似鲍鱼,因此又被称为杏鲍菇。它营养丰富,是口味最好的一种平菇。近年的研究表明刺芹侧耳还具有多种保健活性,因此深受人们的欢迎。但是,两种重要的生理活性物质——多糖和漆酶在刺芹侧耳中的研究报道较少。
原生质体作为遗传操作的重要工具,已经被成功地应用于多种食药用菌的各领域。原生质体无性系变异技术和紫外诱变技术是基于原生质体的两项生物技术,它们在食药用菌中的应用已经为生产中提供了众多优良菌株。但是,这两项技术对于刺芹侧耳的研究报道很少。
随机扩增多态性DNA (RAPD)技术是分子标记的一种,在食用菌种质资源的鉴定、分类、系统发育、遗传多样性、遗传连锁图谱构建以及基因定位、克隆等领域均有良好应用。但是,RAPD技术在刺芹侧耳中的应用,鲜有报道。
基于上述状况,本论文首先对影响刺芹侧耳原生质体制备的因素进行系统优化,旨在建立高效的刺芹侧耳原生质体制备体系；其次,对影响刺芹侧耳原生质体再生条件进行系统研究,以期获得高效的原生质体再生体系；再次,建立刺芹侧耳原生质体无性系变异技术和紫外诱变技术高产多糖菌株和漆酶菌株的筛选体系,以获得高产多糖和漆酶菌株；最后,建立高产菌株的RAPD分子标记体系,以考察菌株间的遗传变异。
结果表明：采用溶壁酶制备刺芹侧耳原生质体的最佳条件为：菌龄、酶解温度、pH、底物浓度、溶壁酶浓度和水解时间分别为6d、30℃、6.0、200mg/ml、1.5%和2.5h,此时刺芹侧耳原生质体的产量为6.34×107个/ml。
刺芹侧耳原生质最佳再生条件为：稳渗剂和稀释剂均为0.6M的甘露醇,培养方式为单层涂布培养,再生培养基为PPR培养基,此时原生质体再生率为0.76%。
原生质体无性系变异技术筛选的高产IPS、EPS、漆酶菌株分别占总无性系菌株的35.7%、39.3%和50%。。IPS、EPS含量及漆酶活力最高的菌株分别是HF15、HF17、HM41菌株,含量或活力分别为127.8±3.2mg/g、324.3±8.3mg/1和5488±101U.min-1.m1-1,分别比亲本HB3菌株高34.8%,42.1%和22.5%。
原生质体紫外诱变技术筛选的高产IPS、EPS、漆酶菌株分别占总诱变株的42.4%、39.4%和15.2%。IPS、EPS含量及漆酶活力最高的是HU30-2、HU50-15、HU50-12菌株,含量或活力分别为114.1±2.7mg/g、360.9±8.0mg/l、5889±108U.ml-1.1-1,分别比亲本HB3菌株高20.4%、58.1%和31.4%,它们的遗传稳定。
确立了邻联甲苯胺法测定刺芹侧耳漆酶活力的最适条件,即波长为620nm、温度为30℃C、pH为4.2、邻联甲苯胺浓度为5mmol/1、反应时间为反应开始至3min。此时,刺芹侧耳漆酶活力比筛选前提高了98%。
采用RAPD分子标记对高产菌株和亲本菌株的遗传差异分析表明：HU50-15菌株变异最大,HM41菌株次之,HF15、HU30-2、HU50-12菌株的变异较小,而HF17菌株未发生RAPD位点的变异。
本文为刺芹侧耳IPS、EPS和漆酶高产菌株的筛选提供了新的方法,不但可以为生产上提供优良的多糖和漆酶高产菌株,而且拓宽了原生质体无性系变异技术和紫外诱变技术研究和应用领域,具有一定的理论意义和现实意义。
Two major types of important bioactive metabolites which were polysaccharides and laccase from P. eryngii had been attracted more and more attentions. Fungi polysaccharides as "biological effects modifier" had very strong nonspecific immunity enhancement effect and antitumor activity through various mechanisms. At present numerous fungi polysaccharides in clinical application had been used for the treatment of diseases, such as tumor, immune deficiency, virus infection diseases and so on. In addition, Fungi polysaccharides also had been developed for functional food additives as they had many other kinds of pharmacological activities such as antioxidative activities, hypoglycemic effect, anti-hyperlipidemic effect and so on. Fungi laccases had important roles on lignin degradation and had extensive substrates so that they had wide application in environmental protection, food, pulp, textile and other industrial fields.
P. eryngii was a kind of rare edible and medicinal fungi which was introduced in the1990s from foreign country. It was also called "xingbaogu" because of its almond-like fragrance and abalone-like flesh. It was quite popular because it was rich in nutrition and delicious best among oyster mushrooms and had all kinds of health cares suggested by recent researches. However, the publications about two important bioactive metaoblites that were polysaccharides and laccase for P. eryngii were less available.
Protoplast was a tool for genetic manipulation which had been successfully applied in varieties of edible and medicinal fungi. Protoplast clonal variation technology and protoplast UV mutagenesis technology were all based on protoplast and had been achieved numerous high-quality and high-quantity strains from edible and medicinal fungi. However, the two technologies had been so rarely applied to P. eryngii.
Random amplified polymorphic DNA (RAPD) was a kind of molecular markers, had been used successfully in genetic and breeding fields such as germplasm identification, classification, phylogenesis, genetic diversity, genetic linkage map construction and gene location and cloning. However, application of RAPD in P. eryngii had been reported less.
Based on the above conditions, firstly, high efficient preparation system of protoplast from P. eryngii was established through systematic optimization of influence factors during protoplast isolation in this paper. Secondly, high efficient regeneration system of protoplast was constructed according to the effect of all factors on regeneration rate. Again, screening systems of high-yield polysaccharides and laccase strains by protoplast clonal variation technology and UV mutagenesis technology were established in order to screen high-yield strains. Finally, RAPD technology system was constructed in order to identify the genetic difference between screened strains and parental strain.
The results showed that the optimum culture days was6d, hydrolytic temperature, pH, substrate concentration, enzyme concentration and reaction time of lywallzyme was30℃,6.0,200mg/ml,1.5%and2.5hours, respectively. Under this condition, the yield of protoplast was up to6.34×107cells/ml.
The best regeneration conditions for the protoplast were followed as:osmotic stabilizer and dilution agent were all0.6M mannitol, regeneration culture mode was single layer and spreading culture, regeneration culture medium was PPR culture medium. Under this condition, protoplast regeneration rate was0.76%.
High-yield IPS, EPS and laccase strains screened by protoplast clonal variation technology accounted for35.7%,39.3%and50%among protoclones strains, respectively. Strain HF15, HF17, HM41showed the highest IPS, EPS content or laccase acitivy of127.8±3.2mg/g,324.3±8.3mg/1and5488±101U.min-1.ml-1which was more34.8%,42.1%and22.5%than parental strain, respectively.
High-yield IPS, EPS and laccase strains screened by protoplast UV mutagenesis technology accounted for42.4%,39.4%and15.2%, respectively. Strain HU30-2, HU50-15, HU50-12showed the highest IPS, EPS content or laccase actitivy of114.1±2.7mg/g,360.9±8.0mg/l,5889+108U.ml-1.1-1, which was more20.4%,58.1%and31.4%than parental strain, respectively. Their yields or activities had genetic stability.
The optimum conditions to measure laccase activity of P. eryngii using ortho-tolidine method were determined:the wavelength was620nm, temperature was30℃, pH was4.2, ortho-tolidine concentration was5mmol/l, reaction time was within3min. Under this condition, laccase activity was highest and98%than before screening.
Genetic differences between high-yield strains and parental strain HB3were analyzed using RAPD technology and the results showed that genetic variation of strain HU50-15comparison with strain HB3was the biggest and HM41was followed. Genetic variation of strain HF15, HU30-12, HU50-12was smaller and variation of strain HF17didn't occur.
New methods were offered to screen high-yield polysaccharides and laccase strains from P. eryngii in this paper and it could not only provide high-yield polysaccharides and laccase strains for production but also broaden application fields of the protoplast clonal variation technology and UV mutagenesis technology and thus it had potential theoretical and realistic significance.
刺芹侧耳(Pleurotus eryngii),是20世纪90年代我国引种成功的一种珍稀食药用真菌,因其据有杏仁的香味,菌肉肥厚似鲍鱼,因此又被称为杏鲍菇。它营养丰富,是口味最好的一种平菇。近年的研究表明刺芹侧耳还具有多种保健活性,因此深受人们的欢迎。但是,两种重要的生理活性物质——多糖和漆酶在刺芹侧耳中的研究报道较少。
原生质体作为遗传操作的重要工具,已经被成功地应用于多种食药用菌的各领域。原生质体无性系变异技术和紫外诱变技术是基于原生质体的两项生物技术,它们在食药用菌中的应用已经为生产中提供了众多优良菌株。但是,这两项技术对于刺芹侧耳的研究报道很少。
随机扩增多态性DNA (RAPD)技术是分子标记的一种,在食用菌种质资源的鉴定、分类、系统发育、遗传多样性、遗传连锁图谱构建以及基因定位、克隆等领域均有良好应用。但是,RAPD技术在刺芹侧耳中的应用,鲜有报道。
基于上述状况,本论文首先对影响刺芹侧耳原生质体制备的因素进行系统优化,旨在建立高效的刺芹侧耳原生质体制备体系；其次,对影响刺芹侧耳原生质体再生条件进行系统研究,以期获得高效的原生质体再生体系；再次,建立刺芹侧耳原生质体无性系变异技术和紫外诱变技术高产多糖菌株和漆酶菌株的筛选体系,以获得高产多糖和漆酶菌株；最后,建立高产菌株的RAPD分子标记体系,以考察菌株间的遗传变异。
结果表明：采用溶壁酶制备刺芹侧耳原生质体的最佳条件为：菌龄、酶解温度、pH、底物浓度、溶壁酶浓度和水解时间分别为6d、30℃、6.0、200mg/ml、1.5%和2.5h,此时刺芹侧耳原生质体的产量为6.34×107个/ml。
刺芹侧耳原生质最佳再生条件为：稳渗剂和稀释剂均为0.6M的甘露醇,培养方式为单层涂布培养,再生培养基为PPR培养基,此时原生质体再生率为0.76%。
原生质体无性系变异技术筛选的高产IPS、EPS、漆酶菌株分别占总无性系菌株的35.7%、39.3%和50%。。IPS、EPS含量及漆酶活力最高的菌株分别是HF15、HF17、HM41菌株,含量或活力分别为127.8±3.2mg/g、324.3±8.3mg/1和5488±101U.min-1.m1-1,分别比亲本HB3菌株高34.8%,42.1%和22.5%。
原生质体紫外诱变技术筛选的高产IPS、EPS、漆酶菌株分别占总诱变株的42.4%、39.4%和15.2%。IPS、EPS含量及漆酶活力最高的是HU30-2、HU50-15、HU50-12菌株,含量或活力分别为114.1±2.7mg/g、360.9±8.0mg/l、5889±108U.ml-1.1-1,分别比亲本HB3菌株高20.4%、58.1%和31.4%,它们的遗传稳定。
确立了邻联甲苯胺法测定刺芹侧耳漆酶活力的最适条件,即波长为620nm、温度为30℃C、pH为4.2、邻联甲苯胺浓度为5mmol/1、反应时间为反应开始至3min。此时,刺芹侧耳漆酶活力比筛选前提高了98%。
采用RAPD分子标记对高产菌株和亲本菌株的遗传差异分析表明：HU50-15菌株变异最大,HM41菌株次之,HF15、HU30-2、HU50-12菌株的变异较小,而HF17菌株未发生RAPD位点的变异。
本文为刺芹侧耳IPS、EPS和漆酶高产菌株的筛选提供了新的方法,不但可以为生产上提供优良的多糖和漆酶高产菌株,而且拓宽了原生质体无性系变异技术和紫外诱变技术研究和应用领域,具有一定的理论意义和现实意义。
Two major types of important bioactive metabolites which were polysaccharides and laccase from P. eryngii had been attracted more and more attentions. Fungi polysaccharides as "biological effects modifier" had very strong nonspecific immunity enhancement effect and antitumor activity through various mechanisms. At present numerous fungi polysaccharides in clinical application had been used for the treatment of diseases, such as tumor, immune deficiency, virus infection diseases and so on. In addition, Fungi polysaccharides also had been developed for functional food additives as they had many other kinds of pharmacological activities such as antioxidative activities, hypoglycemic effect, anti-hyperlipidemic effect and so on. Fungi laccases had important roles on lignin degradation and had extensive substrates so that they had wide application in environmental protection, food, pulp, textile and other industrial fields.
P. eryngii was a kind of rare edible and medicinal fungi which was introduced in the1990s from foreign country. It was also called "xingbaogu" because of its almond-like fragrance and abalone-like flesh. It was quite popular because it was rich in nutrition and delicious best among oyster mushrooms and had all kinds of health cares suggested by recent researches. However, the publications about two important bioactive metaoblites that were polysaccharides and laccase for P. eryngii were less available.
Protoplast was a tool for genetic manipulation which had been successfully applied in varieties of edible and medicinal fungi. Protoplast clonal variation technology and protoplast UV mutagenesis technology were all based on protoplast and had been achieved numerous high-quality and high-quantity strains from edible and medicinal fungi. However, the two technologies had been so rarely applied to P. eryngii.
Random amplified polymorphic DNA (RAPD) was a kind of molecular markers, had been used successfully in genetic and breeding fields such as germplasm identification, classification, phylogenesis, genetic diversity, genetic linkage map construction and gene location and cloning. However, application of RAPD in P. eryngii had been reported less.
Based on the above conditions, firstly, high efficient preparation system of protoplast from P. eryngii was established through systematic optimization of influence factors during protoplast isolation in this paper. Secondly, high efficient regeneration system of protoplast was constructed according to the effect of all factors on regeneration rate. Again, screening systems of high-yield polysaccharides and laccase strains by protoplast clonal variation technology and UV mutagenesis technology were established in order to screen high-yield strains. Finally, RAPD technology system was constructed in order to identify the genetic difference between screened strains and parental strain.
The results showed that the optimum culture days was6d, hydrolytic temperature, pH, substrate concentration, enzyme concentration and reaction time of lywallzyme was30℃,6.0,200mg/ml,1.5%and2.5hours, respectively. Under this condition, the yield of protoplast was up to6.34×107cells/ml.
The best regeneration conditions for the protoplast were followed as:osmotic stabilizer and dilution agent were all0.6M mannitol, regeneration culture mode was single layer and spreading culture, regeneration culture medium was PPR culture medium. Under this condition, protoplast regeneration rate was0.76%.
High-yield IPS, EPS and laccase strains screened by protoplast clonal variation technology accounted for35.7%,39.3%and50%among protoclones strains, respectively. Strain HF15, HF17, HM41showed the highest IPS, EPS content or laccase acitivy of127.8±3.2mg/g,324.3±8.3mg/1and5488±101U.min-1.ml-1which was more34.8%,42.1%and22.5%than parental strain, respectively.
High-yield IPS, EPS and laccase strains screened by protoplast UV mutagenesis technology accounted for42.4%,39.4%and15.2%, respectively. Strain HU30-2, HU50-15, HU50-12showed the highest IPS, EPS content or laccase actitivy of114.1±2.7mg/g,360.9±8.0mg/l,5889+108U.ml-1.1-1, which was more20.4%,58.1%and31.4%than parental strain, respectively. Their yields or activities had genetic stability.
The optimum conditions to measure laccase activity of P. eryngii using ortho-tolidine method were determined:the wavelength was620nm, temperature was30℃, pH was4.2, ortho-tolidine concentration was5mmol/l, reaction time was within3min. Under this condition, laccase activity was highest and98%than before screening.
Genetic differences between high-yield strains and parental strain HB3were analyzed using RAPD technology and the results showed that genetic variation of strain HU50-15comparison with strain HB3was the biggest and HM41was followed. Genetic variation of strain HF15, HU30-12, HU50-12was smaller and variation of strain HF17didn't occur.
New methods were offered to screen high-yield polysaccharides and laccase strains from P. eryngii in this paper and it could not only provide high-yield polysaccharides and laccase strains for production but also broaden application fields of the protoplast clonal variation technology and UV mutagenesis technology and thus it had potential theoretical and realistic significance.
引文
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