斜卧青霉β-葡萄糖苷酶的性质和功能研究及基因表达谱分析
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摘要
木质纤维素是地球上储量最为丰富的可再生资源。在矿物燃料资源日趋枯竭、环境日趋恶化的严峻形势下,利用廉价的木质纤维素生产液体燃料和大宗化学品,可以有效地缓解能源和环境危机,促进全球经济的可持续发展。自然界中存在多种高效木质纤维素生物降解系统,利用微生物产生的木质纤维素水解酶来降解木质纤维素是纤维素利用的一种有效途径。丝状真菌在自然生态环境木质纤维素的降解中发挥着非常重要的作用,对其木质纤维素酶酶系组成、催化机理和调控机制的研究具有重要的理论意义与应用价值。
斜卧青霉114-2能分泌完全的纤维素酶酶系,与里氏木霉相比具有较高的p-葡萄糖苷酶及半纤维素酶活性,这对于木质纤维素材料的水解是非常有利的。但是较高的p-葡萄糖苷酶活力使得纤维素底物迅速水解为葡萄糖,不利于诱导物(纤维二糖)的积累。因此从纤维素酶的合成调控的角度考虑,胞外过多的p-葡萄糖苷酶的存在对纤维素酶的诱导是不利的。由此可见,β-葡萄糖苷酶作为木质纤维素降解酶系的重要组分,对于其它酶组分的合成有一定的调控作用。
目前,对斜卧青霉基因组的测序工作已经完成,对其纤维素酶合成调控的研究也已取得一定的进展。但是由于其纤维素酶酶系复杂,涉及的调控层次和调控途径众多,传统的针对单基因的研究方法很难揭示木质纤维素酶合成调控的整体网络。而转录组学研究可以揭示不同菌株在不同培养条件下全基因组范围的基因转录差异,有助于发现与木质纤维素酶合成调控相关的新基因,完善对合成调控网络的认识,找到工程改造的新靶点。本论文的主要研究结果如下：
1.胞外主要p-葡萄糖苷酶BGL1的性质及其在木质纤维素酶合成调控中的功能研究
在斜卧青霉114-2胞外粗酶液中分离纯化到一个p-葡萄糖苷酶,质谱鉴定其为胞外主要的β-葡萄糖苷酶BGL1,具有很高的比活力、温度和pH稳定性及较高的底物亲和力。对里氏木霉QM9414胞外粗酶的糖化能力具有很强的促进作用,与商业酶制剂的效果相当。
在斜卧青霉114-2中对BGL1进行了敲除,发现BGL1缺失后突变株在葡萄糖、麦麸及纤维二糖平板上的生长和表型都未受到影响。同时,在产酶条件下其胞外蛋白的分泌能力及CBH、EG和木聚糖酶活力也没有发生明显变化。与胞外酶活力的结果一致,在纤维素条件下突变株中主要的木质纤维素酶基因的转录水平与野生菌株相比也没有发生明显的变化。对野生株和突变株在纤维素条件下胞外的纤维寡糖浓度进行测定,发现突变株胞外的纤维寡糖大量积累。但是纤维寡糖的积累却没有诱导木质纤维素酶基因的表达上调。同时当以纤维寡糖为唯一碳源进行培养时,也不能诱导突变株木质纤维素酶基因的表达上调。我们推测,斜卧青霉不能直接感受胞外的寡糖诱导信号来诱导木质纤维素酶基因的表达,纤维寡糖只有进入胞内才能行使诱导作用。通过测定纤维素诱导条件下胞内的纤维寡糖浓度,发现突变株胞内纤维二糖的浓度与野生株相似,与木质纤维素酶基因表达的趋势是一致的。而在胞外添加纤维二糖时,突变株胞内纤维二糖的浓度较野生株有所提高,同时木质纤维素酶基因的表达上调。这说明胞内纤维二糖的浓度是影响木质纤维素酶基因表达水平的关键。而BGL1的缺失之所以没有造成胞内纤维二糖的积累,推测是因为胞内β-葡萄糖苷酶的存在。2.胞内主要β-葡萄糖苷酶BGL2的性质及其在木质纤维素酶合成调控中的功能
研究将斜卧青霉基因组中6个推测的胞内p-葡萄糖苷酶分别在野生菌株114-2中进行了基因敲除,发现在麦麸、葡萄糖和纤维二糖平板上各突变株的表型与野生菌株相比均没有明显变化。只有在表达量最高的BGL2缺失后,突变株在产酶培养基中的胞外蛋白产量和木质纤维素酶活力才表现出大幅度提高。SDS-PAGE结果显示,其胞外蛋白浓度的升高并不是整体分泌量增加引起的,而是木质纤维素酶比例的特异性提高造成的。胞外酶液EG和木聚糖酶比活力的相应提高也证实也这一结果。与蛋白水平变化一致,敲除株中的木质纤维素酶基凶egl、cbhl和xyn10的表达水平在纤维素条件下均明显上调。同时发现其胞内纤维二糖浓度较野生株有所升高。这些结果显示,是由于BGL2的缺失使得敲除株丧失大部分胞内p-葡萄糖苷酶活力,致使胞内纤维二糖积累,从而诱导了木质纤维素酶基因的高水平表达。将BGL2在大肠杆菌中异源表达,对重组酶的酶学性质进行分析。结果发现,BGL2对纤维二糖、龙胆二糖、槐糖和纤维寡糖都有很强的水解活性。这就证实了野生菌株中BGL2的存在会迅速水解胞内的纤维寡糖,不利于诱导物的积累,从而限制木质纤维素酶基因的表达水平。
3.p-葡萄糖苷酶双敲除株中木质纤维素酶诱导机制的研究
构建了BGL1和BGL2双缺失菌株△bgllΔbgl2,发现其在麦麸、葡萄糖和纤维二糖平板上的生长及形态没有发生明显变化,胞外木质纤维素酶活力相对于野生菌株均明显提高,但是低于△bgl2的酶活力水平。与蛋白水平变化一致,在纤维素条件下敲除株中主要的木质纤维素酶基因的转录水平较野生株明显提高,但是也低于Δbg12中的表达水平。测定纤维素条件下的胞内纤维二糖浓度,显示其浓度变化趋势与木质纤维素酶基因的转录水平一致。我们猜测ΔbgllΔbgl2不能在胞内积累更多的纤维二糖的原因可能是,斜卧青霉在β-葡萄糖苷酶活性缺失时,会激发其它的代谢途径来利用纤维二糖。用潜在诱导物槐糖和龙胆二糖作为唯一碳源进行诱导,发现纤维素酶基因的表达水平并没有发生明显上调。这说明槐糖和龙胆二糖不是斜卧青霉纤维素酶基因的诱导物,而且在斜卧青霉中由纤维二糖介导的诱导作用并不依赖于通过转糖基作用牛成槐糖或龙胆二糖的过程。4.斜卧青霉野生菌株114-2及突变株JU-A10-T的差异表达谱分析
利用数字化表达谱分析技术对斜卧青霉不同菌株在不同条件下的转录组进行了研究。分别比较了野生菌株114-2及突变株JU-A10-T在不同条件下及在相同条件下114-2与JU-A10-T之间的表达差异基因,分析了与木质纤维素酶基因共转录的差异表达基因,筛选到一系列可能与木质纤维素酶基因表达上调有关的候选基因,但其功能还有待验证。研究发现,JU-A10-T中次级代谢相关、淀粉酶及蛋白酶等基因的转录水平相对于野生菌株114-2均明显降低。这可能通过缓解蛋白质合成负担、减少不必要的能量消耗等机制部分导致了JU-A10-T中纤维素酶基因表达水平的提高。
Lignocellulosic biomass is the largest reserves of renewable resource on the earth. With the intensification of fossil fuels shortage and environment pollution, effective use of these low-cost resource to produce biofuel and biochemical is an efficient way to relieve the energy crisis, protect environment and maintain sustainable economic development. There are many lignocellulose degradation systems in nature, in which conversion by enzyme systems from microorganism was one of the most efficient pathways. Fungi play an important role in the environmental lignocellulose degradation. The researches on the composition of their lignocellulolytic enzyme system and the regulation of enzyme expression are very important for improvement of cellulase production and enzyme system composition.
The filamentous fungus Penicillium decumbens114-2can produce a balanced extracellular lignocellulolytic enzyme system with more β-glucosidase, which is very efficient for lignocellulose degradation. However, the plenty of β-glucosidases can hydrolyze the oligosaccharides to glucose immediately, which restricts the accumulation of inducers and the induction of cellulase genes. It shows that, as an important component of lignocellulolytic enzyme system, β-glucosidase can also participate in the regulation of cellulase gene expression by regulating the concentration of inducer.
There have been several researches about the regulation mechanisms for lignocellulolytic enzymes production in P. decumbens. However, traditional methods studying on single-gene is hard to unravel the holistic regulation mechanisms for lignocellulolytic enzymes production due to the complexity of cellulase enzyme system and multifarious of regulatory levels and pathways. Transcriptomics, which can reveal the genome-wide differences in gene transcription of different strains under different culture conditions, is very useful in perfecting the holistic network of regulation mechanisms for lignocellulolytic enzymes production.
The main results of the thesis are as follows:
1. Characterization of the major extracellular β-glucosidase BGL1and its role in regulation of lignocellulolytic enzymes
A β-glucosidase was purified from extracellular enzymes of P. decumbens114-2, which was identified the major extracellular β-glucosidase BGL1by MS/MS. The properties of BGL1, including its pH and temperature optima, the high affinity to substrates and high specific activity, make it has great potential to be utilized as supplementation in conversion of corncob residue and other lignocellulosic biomass into simple sugars. And it showed similar ability in boosting enzymatic degradation of substrate with the P-glucosidase from commercial enzymes NS-50010.
After deletion of bgll, the phenotypes, including colony radial, growth rate, conidia formation or conidia color, were not affected in Abgll. When incubated in CW (cellulose+wheat bran) medium, the production of extracellular proteins (except for BGL1) was almost the same as the wild type strain. In accordance with this, expression levels of the major extracellular lignocellulolytic enzyme genes egl, cbhl and xyn10were similar with the parent strain on1%MCC in24h, although the extracellular oligosaccharides were accumulated during the incubation. Meanwhile, addition of oligosaccharides in the medium could not induce the higher expression of cellulase genes. When the intracellular cellobiose was determined on MCC (microcrystalline cellulose) medium, no more cellobiose was accumulated in Abgll, which might be due to the existence of intracellular β-glucosidase. The expressions of detected genes were all up-regulated after cultivation with0.02%cellobiose for4h, while the intracellular cellobiose concentration was in synchronous with gene repressions. We assumed that the oligosaccharides should be transported into the cell to trigger the induction, and the intracellular cellobiose concentration was important for the induction degree.
2. Characterization of the major intracellular β-glucosidase BGL1and its role in regulation of lignocellulolytic enzymes
The six intracellular p-glucosidases were deleted in P. decumbens114-2 respectively. As the major intracellular β-glucosidase, the absence of BGL2improved the extracellular FPA, EG, CBH and xylanase activities dramatically on CW medium compare with the wild type strain. While deletion of other β-glucosidases barely make any difference. SDS-PAGE analysis suggested that the secretion improvement of the extracellular proteins was not general and only some of proteins increased dramatically in Abgl2, which were probably lignocellulolytic enzymes. Meanwhile the calculated specific EG and xylanase activities of Abgl2broth also confirmed the conclusion. As a constitutive enzyme, the production of extracellular β-glucosidase was not promoted synchronously with those of cellulose-induced enzymes. In accordance with the enzymes production, the transcription levels of egl, cbhl, xynlOA were improved dramatically on0.02%cellobiose and1%MCC in Abgl2. When the intracellular cellodextrins of P. decumbens114-2and Abgl2were analyzed, greater amounts of cellobiose was determined in the cytoplasm of Abgl2than that of114-2. To explore the mechanism of the improved production of lignocellulolytic enzymes by deficiency of BGL2, bgl2was expressed in E. coli BL21(DE3) and the characters of purified rBGL2were analyzed. rBGL2showed efficient hydrolytic activities toward cellodextrins with degree of polymerization from2to4. Besides cellodextrins with the glycosidic bond type of β-1,4, rBGL2could also hydrolyze sophorose (β-1,2bond) and gentiobiose (β-1,6bond). The results strongly supported that the improved induction of lignocellulolytic enzymes expression in Abgl2by cellobiose and cellulose might be mediated by the intracellular accumulation of cellobiose.
3. Preliminary study of the cellulase induction mechanism of P. decumbens in double deletion strain of P-glucosidase genes
The mutant lacking both bglland bgl2was constructed. The mutant AbgllΔbgl2lost nearly all of the extra-β-glucosidase and most of intracellular β-glucosidase activities. However, AbgllΔbgl2could survive on the medium with cellobiose as the sole carbon source normally. The production of extracellular lignocellulolytic enzymes was not further enhanced compared with that of Abgl2as expected when incubated in CW medium. Transcriptional analysis of egl, cbhl and xyn10also revealed that simultaneously deletion of bgll and bgl2did not cause the further improvement of lignocellulolytic genes expression when incubated either on cellobiose or MCC. In accord with this, there was less cellobiose determined in the cytoplasm of AbgllΔbgl2compared with that of Abgl2, which probably be the reason resulting in the less efficient expression of lignocellulolytic genes. The results suggested that there seemed to be some other pathways activated to utilize cellobiose in the absence of main β-glucosidases, which caused the less accumulation of intracellular cellobiose and lower production of lignocellulolytic enzymes in AbgllAbgl2. The transcription levels of egl and cbhl were analyzed when induced by two potential inducers sophorose and gentiobiose in mutant AbgllAbgl2and no inductive effect was observed. The results suggested that despite acting as cellulase inducers in some other fungi, the two disaccharides did not participate in the induction of cellulase expression in P. decumbens. In other words, the induction mediated by cellobiose and cellulose was unrelated to the formation, if any, of sophorose or gentiobiose from cellobiose in P. decumbens.
4. Transcriptional analysis of P. decumbens wild type strain114-2and mutant JU-A10-T
Transcriptome of P. decumbens114-2and JU-A10-T on different carbon sources were analyzed using digital gene expression tag profiling (DGE) technology. The differential expression genes between114-2and JU-A10-T, as well as the differential expression genes on different carbon sources, were analyzed by Gene Ontology functional enrichment. The genes co-regulated with lignocellulolytic enzyme genes were worth studying, which might be related to the up-regulation of lignocellulolytic enzyme genes. The down-regulated genes in JU-A10-T relative to114-2were involved in organics transport, secondary metabolism and synthesizing amylases and proteases, which might contribute to the up-regulation of lignocellulolytic enzyme genes by alleviating burdens of protein synthesis and energy consumption.
斜卧青霉114-2能分泌完全的纤维素酶酶系,与里氏木霉相比具有较高的p-葡萄糖苷酶及半纤维素酶活性,这对于木质纤维素材料的水解是非常有利的。但是较高的p-葡萄糖苷酶活力使得纤维素底物迅速水解为葡萄糖,不利于诱导物(纤维二糖)的积累。因此从纤维素酶的合成调控的角度考虑,胞外过多的p-葡萄糖苷酶的存在对纤维素酶的诱导是不利的。由此可见,β-葡萄糖苷酶作为木质纤维素降解酶系的重要组分,对于其它酶组分的合成有一定的调控作用。
目前,对斜卧青霉基因组的测序工作已经完成,对其纤维素酶合成调控的研究也已取得一定的进展。但是由于其纤维素酶酶系复杂,涉及的调控层次和调控途径众多,传统的针对单基因的研究方法很难揭示木质纤维素酶合成调控的整体网络。而转录组学研究可以揭示不同菌株在不同培养条件下全基因组范围的基因转录差异,有助于发现与木质纤维素酶合成调控相关的新基因,完善对合成调控网络的认识,找到工程改造的新靶点。本论文的主要研究结果如下：
1.胞外主要p-葡萄糖苷酶BGL1的性质及其在木质纤维素酶合成调控中的功能研究
在斜卧青霉114-2胞外粗酶液中分离纯化到一个p-葡萄糖苷酶,质谱鉴定其为胞外主要的β-葡萄糖苷酶BGL1,具有很高的比活力、温度和pH稳定性及较高的底物亲和力。对里氏木霉QM9414胞外粗酶的糖化能力具有很强的促进作用,与商业酶制剂的效果相当。
在斜卧青霉114-2中对BGL1进行了敲除,发现BGL1缺失后突变株在葡萄糖、麦麸及纤维二糖平板上的生长和表型都未受到影响。同时,在产酶条件下其胞外蛋白的分泌能力及CBH、EG和木聚糖酶活力也没有发生明显变化。与胞外酶活力的结果一致,在纤维素条件下突变株中主要的木质纤维素酶基因的转录水平与野生菌株相比也没有发生明显的变化。对野生株和突变株在纤维素条件下胞外的纤维寡糖浓度进行测定,发现突变株胞外的纤维寡糖大量积累。但是纤维寡糖的积累却没有诱导木质纤维素酶基因的表达上调。同时当以纤维寡糖为唯一碳源进行培养时,也不能诱导突变株木质纤维素酶基因的表达上调。我们推测,斜卧青霉不能直接感受胞外的寡糖诱导信号来诱导木质纤维素酶基因的表达,纤维寡糖只有进入胞内才能行使诱导作用。通过测定纤维素诱导条件下胞内的纤维寡糖浓度,发现突变株胞内纤维二糖的浓度与野生株相似,与木质纤维素酶基因表达的趋势是一致的。而在胞外添加纤维二糖时,突变株胞内纤维二糖的浓度较野生株有所提高,同时木质纤维素酶基因的表达上调。这说明胞内纤维二糖的浓度是影响木质纤维素酶基因表达水平的关键。而BGL1的缺失之所以没有造成胞内纤维二糖的积累,推测是因为胞内β-葡萄糖苷酶的存在。2.胞内主要β-葡萄糖苷酶BGL2的性质及其在木质纤维素酶合成调控中的功能
研究将斜卧青霉基因组中6个推测的胞内p-葡萄糖苷酶分别在野生菌株114-2中进行了基因敲除,发现在麦麸、葡萄糖和纤维二糖平板上各突变株的表型与野生菌株相比均没有明显变化。只有在表达量最高的BGL2缺失后,突变株在产酶培养基中的胞外蛋白产量和木质纤维素酶活力才表现出大幅度提高。SDS-PAGE结果显示,其胞外蛋白浓度的升高并不是整体分泌量增加引起的,而是木质纤维素酶比例的特异性提高造成的。胞外酶液EG和木聚糖酶比活力的相应提高也证实也这一结果。与蛋白水平变化一致,敲除株中的木质纤维素酶基凶egl、cbhl和xyn10的表达水平在纤维素条件下均明显上调。同时发现其胞内纤维二糖浓度较野生株有所升高。这些结果显示,是由于BGL2的缺失使得敲除株丧失大部分胞内p-葡萄糖苷酶活力,致使胞内纤维二糖积累,从而诱导了木质纤维素酶基因的高水平表达。将BGL2在大肠杆菌中异源表达,对重组酶的酶学性质进行分析。结果发现,BGL2对纤维二糖、龙胆二糖、槐糖和纤维寡糖都有很强的水解活性。这就证实了野生菌株中BGL2的存在会迅速水解胞内的纤维寡糖,不利于诱导物的积累,从而限制木质纤维素酶基因的表达水平。
3.p-葡萄糖苷酶双敲除株中木质纤维素酶诱导机制的研究
构建了BGL1和BGL2双缺失菌株△bgllΔbgl2,发现其在麦麸、葡萄糖和纤维二糖平板上的生长及形态没有发生明显变化,胞外木质纤维素酶活力相对于野生菌株均明显提高,但是低于△bgl2的酶活力水平。与蛋白水平变化一致,在纤维素条件下敲除株中主要的木质纤维素酶基因的转录水平较野生株明显提高,但是也低于Δbg12中的表达水平。测定纤维素条件下的胞内纤维二糖浓度,显示其浓度变化趋势与木质纤维素酶基因的转录水平一致。我们猜测ΔbgllΔbgl2不能在胞内积累更多的纤维二糖的原因可能是,斜卧青霉在β-葡萄糖苷酶活性缺失时,会激发其它的代谢途径来利用纤维二糖。用潜在诱导物槐糖和龙胆二糖作为唯一碳源进行诱导,发现纤维素酶基因的表达水平并没有发生明显上调。这说明槐糖和龙胆二糖不是斜卧青霉纤维素酶基因的诱导物,而且在斜卧青霉中由纤维二糖介导的诱导作用并不依赖于通过转糖基作用牛成槐糖或龙胆二糖的过程。4.斜卧青霉野生菌株114-2及突变株JU-A10-T的差异表达谱分析
利用数字化表达谱分析技术对斜卧青霉不同菌株在不同条件下的转录组进行了研究。分别比较了野生菌株114-2及突变株JU-A10-T在不同条件下及在相同条件下114-2与JU-A10-T之间的表达差异基因,分析了与木质纤维素酶基因共转录的差异表达基因,筛选到一系列可能与木质纤维素酶基因表达上调有关的候选基因,但其功能还有待验证。研究发现,JU-A10-T中次级代谢相关、淀粉酶及蛋白酶等基因的转录水平相对于野生菌株114-2均明显降低。这可能通过缓解蛋白质合成负担、减少不必要的能量消耗等机制部分导致了JU-A10-T中纤维素酶基因表达水平的提高。
Lignocellulosic biomass is the largest reserves of renewable resource on the earth. With the intensification of fossil fuels shortage and environment pollution, effective use of these low-cost resource to produce biofuel and biochemical is an efficient way to relieve the energy crisis, protect environment and maintain sustainable economic development. There are many lignocellulose degradation systems in nature, in which conversion by enzyme systems from microorganism was one of the most efficient pathways. Fungi play an important role in the environmental lignocellulose degradation. The researches on the composition of their lignocellulolytic enzyme system and the regulation of enzyme expression are very important for improvement of cellulase production and enzyme system composition.
The filamentous fungus Penicillium decumbens114-2can produce a balanced extracellular lignocellulolytic enzyme system with more β-glucosidase, which is very efficient for lignocellulose degradation. However, the plenty of β-glucosidases can hydrolyze the oligosaccharides to glucose immediately, which restricts the accumulation of inducers and the induction of cellulase genes. It shows that, as an important component of lignocellulolytic enzyme system, β-glucosidase can also participate in the regulation of cellulase gene expression by regulating the concentration of inducer.
There have been several researches about the regulation mechanisms for lignocellulolytic enzymes production in P. decumbens. However, traditional methods studying on single-gene is hard to unravel the holistic regulation mechanisms for lignocellulolytic enzymes production due to the complexity of cellulase enzyme system and multifarious of regulatory levels and pathways. Transcriptomics, which can reveal the genome-wide differences in gene transcription of different strains under different culture conditions, is very useful in perfecting the holistic network of regulation mechanisms for lignocellulolytic enzymes production.
The main results of the thesis are as follows:
1. Characterization of the major extracellular β-glucosidase BGL1and its role in regulation of lignocellulolytic enzymes
A β-glucosidase was purified from extracellular enzymes of P. decumbens114-2, which was identified the major extracellular β-glucosidase BGL1by MS/MS. The properties of BGL1, including its pH and temperature optima, the high affinity to substrates and high specific activity, make it has great potential to be utilized as supplementation in conversion of corncob residue and other lignocellulosic biomass into simple sugars. And it showed similar ability in boosting enzymatic degradation of substrate with the P-glucosidase from commercial enzymes NS-50010.
After deletion of bgll, the phenotypes, including colony radial, growth rate, conidia formation or conidia color, were not affected in Abgll. When incubated in CW (cellulose+wheat bran) medium, the production of extracellular proteins (except for BGL1) was almost the same as the wild type strain. In accordance with this, expression levels of the major extracellular lignocellulolytic enzyme genes egl, cbhl and xyn10were similar with the parent strain on1%MCC in24h, although the extracellular oligosaccharides were accumulated during the incubation. Meanwhile, addition of oligosaccharides in the medium could not induce the higher expression of cellulase genes. When the intracellular cellobiose was determined on MCC (microcrystalline cellulose) medium, no more cellobiose was accumulated in Abgll, which might be due to the existence of intracellular β-glucosidase. The expressions of detected genes were all up-regulated after cultivation with0.02%cellobiose for4h, while the intracellular cellobiose concentration was in synchronous with gene repressions. We assumed that the oligosaccharides should be transported into the cell to trigger the induction, and the intracellular cellobiose concentration was important for the induction degree.
2. Characterization of the major intracellular β-glucosidase BGL1and its role in regulation of lignocellulolytic enzymes
The six intracellular p-glucosidases were deleted in P. decumbens114-2 respectively. As the major intracellular β-glucosidase, the absence of BGL2improved the extracellular FPA, EG, CBH and xylanase activities dramatically on CW medium compare with the wild type strain. While deletion of other β-glucosidases barely make any difference. SDS-PAGE analysis suggested that the secretion improvement of the extracellular proteins was not general and only some of proteins increased dramatically in Abgl2, which were probably lignocellulolytic enzymes. Meanwhile the calculated specific EG and xylanase activities of Abgl2broth also confirmed the conclusion. As a constitutive enzyme, the production of extracellular β-glucosidase was not promoted synchronously with those of cellulose-induced enzymes. In accordance with the enzymes production, the transcription levels of egl, cbhl, xynlOA were improved dramatically on0.02%cellobiose and1%MCC in Abgl2. When the intracellular cellodextrins of P. decumbens114-2and Abgl2were analyzed, greater amounts of cellobiose was determined in the cytoplasm of Abgl2than that of114-2. To explore the mechanism of the improved production of lignocellulolytic enzymes by deficiency of BGL2, bgl2was expressed in E. coli BL21(DE3) and the characters of purified rBGL2were analyzed. rBGL2showed efficient hydrolytic activities toward cellodextrins with degree of polymerization from2to4. Besides cellodextrins with the glycosidic bond type of β-1,4, rBGL2could also hydrolyze sophorose (β-1,2bond) and gentiobiose (β-1,6bond). The results strongly supported that the improved induction of lignocellulolytic enzymes expression in Abgl2by cellobiose and cellulose might be mediated by the intracellular accumulation of cellobiose.
3. Preliminary study of the cellulase induction mechanism of P. decumbens in double deletion strain of P-glucosidase genes
The mutant lacking both bglland bgl2was constructed. The mutant AbgllΔbgl2lost nearly all of the extra-β-glucosidase and most of intracellular β-glucosidase activities. However, AbgllΔbgl2could survive on the medium with cellobiose as the sole carbon source normally. The production of extracellular lignocellulolytic enzymes was not further enhanced compared with that of Abgl2as expected when incubated in CW medium. Transcriptional analysis of egl, cbhl and xyn10also revealed that simultaneously deletion of bgll and bgl2did not cause the further improvement of lignocellulolytic genes expression when incubated either on cellobiose or MCC. In accord with this, there was less cellobiose determined in the cytoplasm of AbgllΔbgl2compared with that of Abgl2, which probably be the reason resulting in the less efficient expression of lignocellulolytic genes. The results suggested that there seemed to be some other pathways activated to utilize cellobiose in the absence of main β-glucosidases, which caused the less accumulation of intracellular cellobiose and lower production of lignocellulolytic enzymes in AbgllAbgl2. The transcription levels of egl and cbhl were analyzed when induced by two potential inducers sophorose and gentiobiose in mutant AbgllAbgl2and no inductive effect was observed. The results suggested that despite acting as cellulase inducers in some other fungi, the two disaccharides did not participate in the induction of cellulase expression in P. decumbens. In other words, the induction mediated by cellobiose and cellulose was unrelated to the formation, if any, of sophorose or gentiobiose from cellobiose in P. decumbens.
4. Transcriptional analysis of P. decumbens wild type strain114-2and mutant JU-A10-T
Transcriptome of P. decumbens114-2and JU-A10-T on different carbon sources were analyzed using digital gene expression tag profiling (DGE) technology. The differential expression genes between114-2and JU-A10-T, as well as the differential expression genes on different carbon sources, were analyzed by Gene Ontology functional enrichment. The genes co-regulated with lignocellulolytic enzyme genes were worth studying, which might be related to the up-regulation of lignocellulolytic enzyme genes. The down-regulated genes in JU-A10-T relative to114-2were involved in organics transport, secondary metabolism and synthesizing amylases and proteases, which might contribute to the up-regulation of lignocellulolytic enzyme genes by alleviating burdens of protein synthesis and energy consumption.
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