琼脂降解相关酶类及酶法制备新琼寡糖的研究
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摘要
海洋占地球总面积的70%以上,蕴藏着大量的生物资源,其中海藻资源尤为丰富。从海藻中提取琼脂的工艺都是建立在对海藻原料进行碱处理→水洗→酸处理、漂白→水洗的基础上来进行的,不仅造成了严重的环境污染,而且消耗了大量的水资源。我国出口的琼脂产品多以低附加值的琼脂胶、琼脂粉为主；经国外精细加工后的琼脂寡糖又以较高的价格在国内销售。因此,对现行的琼脂提取工艺进行改进,以期将降解琼脂的相关酶类应用于琼脂生产过程中,利用酶法制备高附加值的琼脂寡糖,上述问题就可迎刃而解,在获得最大经济效益的同时尽可能地避免环境的污染,从而推动这一行业得到更快更好的发展。
本研究,从厦门红树林泥样中筛选到一株硫酸酯酶产生菌S9,通过16SrRNA基因序列的同源性分析,确定该菌株为假单胞菌属。对其全基因组进行测序分析发现,Pseudomonas sp. S9基因组由一个环状染色体构成,总长4.7M,GC%含量为56.52%,有2个rRNA操纵元,56个tRNA编码基因对应全部20种氨基酸。预测共编码4665个ORF。KEGG.COGs分析表明S9菌株中有38.3%的功能基因与代谢相关,该菌株具有较为完整的糖代谢,脂肪酸代谢以及氨基酸代谢等基础代谢途径。另外,在Pseudomonas sp. S9基因组中发现了大量ABCtansporter编码基因和细菌耐高浓度二价重金属离子Me2+的基因簇。这些基因的大量存在可能都与Pseudomonas sp. S9菌株适应红树林复杂生存环境相关。
整个Pseudomonas sp. S9基因组中存在7个与芳香族硫酸酯酶相关的蛋白。其中6个序列完整的芳香族硫酸酯酶都具有CX[PIA]XRX的保守结构域。4个Pseudomonas sp. S9的芳香族硫酸酯具有信号肽,表明这些芳香族硫酸酯不同于其它的Cry型芳香族硫酸酯酶,它们具有分泌到细胞外的潜力。首次报道了具有分泌功能的Cry型芳香族硫酸酯酶。
异源表达了菌株Pseudomonas sp. S9中的芳香族硫酸酯酶,对酶降解龙须菜硫酸根基团进行了相关功能研究。并且,通过对S9菌株全基因组文库的构建,获得了其中能够降解SDS的烷基型硫酸酯酶编码基因sdsAP,大肠杆菌中异源表达后的SdsAP具有很好的温度稳定性,扩展了烷基型硫酸酯酶的酶资源。
从海藻浒苔中筛选到一株高产琼脂酶的菌株LQ48,经鉴定LQ48为噬琼脂菌属淡黄色噬琼脂菌Agarivorans gilvus中的一个新菌株。克隆和重组表达了菌株Agarivorans sp. LQ48的GH16家族β-琼脂酶基因agaA,经过酶学性质分析,其具有广范围的pH稳定性。在琼脂酶AgaA研究基础上,通过LQ48菌株粗琼脂酶降解琼脂糖制备出不同聚合度的新琼寡糖混合物,利用分子筛成功地将新琼二糖NA2和新琼四糖NA4进行分离纯化。体外抗氧化性研究表明酶法制备的NA2和NA4均具有一定清除DPPH自由基、清除超氧阴离子自由基和清除羟基自由基的能力。
本研究的完成,为现行琼脂提取工艺的改进以及探索酶法生产高附加值琼脂产品的研究奠定理论基础。
The oceans are the Earth's largest ecosystem, covering 70% of our planet and providing goods and services for the majority of the world's population, particularly the algae. The extraction of agar is currently based on the procedure of alkali treatment, water washing, acid treatment, bleaching and water washing on raw algae materials, which not only cause serious environmental pollution, but also consumes vast quantities of water resources. On the other hand, the exported agar-relevant products are mainly low value-added agar, while processed agarose and agar oligosaccharides are imported with higher prices in China. Therefore, it is essential to improve the current process of extraction by applying the enzymatic treatment to degrade the sulfates of algae and produce the high value-added agar oligosaccharides, which will help to prevent environmental pollution and to gain the maximum economic benefits, thus promoting a faster and better development of the agar industry.
In this study, we isolated a sulfatase-producing bacterium S9 from the mangrove soil samples.16S rRNA analysis indicated that it belonged to genus Pseudomonas. The genome analysis suggests that the genome of Pseudomonas sp. S9 is consisted of a circular DNA. The draft genome excluding the gaps has a total of 4,796,832 bases comprising 4,665 predicted open reading frames (ORFs), with a G+C content of 56.52%. There are two rRNA operons and 56 predicted tRNA genes. In addition, the genome of Pseudomonas sp. S9 encodes amount of ABC tansporter genes and heavy metal tolerance genes. These are likely to bear systems to cope with toxic mangrove environment. According to the KEGG and COGs analysis, Pseudomonas sp. S9 is predicted that 38.3% function genes are related to the metabolism function. Moreover, it possesses the complete pathway of carbohydrate metabolism, lipid metabolism and amino acid metabolism. Additional, there are amounts of ABC transporter genes and heavy metal tolerance genes. These suggest Pseudomonas sp. S9 can cope with toxic metals in mangrove environment.
In S9 genome, seven predicted ORFs are annotated as arylsulfatases, six of whose sequences are complete. All these complete coding sequences have the CX[P/A]XRX motif, a characteristic of the cysteine-type sulfatases. Using SignalP to search for the possible signal peptides in cysteine-type sulfatases, we find four of them carrying a signal peptide which is accordant with that the arylsulfatases are exited in extracellular of Pseudomonas sp. S9. It is the first report of the extracellular cysteine-type sulfatases.
Meanwhile, arylsulfatase genes were cloned from strain S9 and expressed in E. coli BL21, and the recombinant enzyme was characterized to degrade the sulfate of polysaccharide from Gracilaria Lemaneiformis. That would provide a foundation for enzymatic degradation instead alkaline degradation in producing agar. Moreover, a novel alkylsulfatase gene, sdsAP, was cloned from a newly isolated bacterium Pseudomonas sp. S9 and expressed in Escherichia coli. The noticeable thermal stability makes SdsAP an ideal candidate for the application in disposing SDS-containing waste.
We described the isolation and identification of a new agarolytic bacterium Agarivorans sp. LQ48 from marine algae, and the cloning and expression of a novel endo-typeβ-agarase gene, agaA, in Escherichia coli. We also explored the characteristics of the purified recombinant agarase. On account of its noticeable pH stability, the enzyme can be utilized extensively in many kinds of industries. Then a series of neoagaroligosaccharides with different degree of polymerization were prepared by the crude agarases of strain LQ48. Neoagarobiose and neoagarotetraose were purified by Sephadex G-15 and identified by TLC. Furthermore, their antioxidant activity was investigated by various antioxidant assay in vitro systems, including DPPH radical scavenging, superoxide anion radical scavenging, hydrogen radical scavenging. The results indicated that neoagarobiose and neoagarotetraose producing by enzyme exhibited the antioxidant activities.
This study is the first step of improving the technique of agar production and exploring the methods of producing the neoagaroligosaccharides by agarases.
本研究,从厦门红树林泥样中筛选到一株硫酸酯酶产生菌S9,通过16SrRNA基因序列的同源性分析,确定该菌株为假单胞菌属。对其全基因组进行测序分析发现,Pseudomonas sp. S9基因组由一个环状染色体构成,总长4.7M,GC%含量为56.52%,有2个rRNA操纵元,56个tRNA编码基因对应全部20种氨基酸。预测共编码4665个ORF。KEGG.COGs分析表明S9菌株中有38.3%的功能基因与代谢相关,该菌株具有较为完整的糖代谢,脂肪酸代谢以及氨基酸代谢等基础代谢途径。另外,在Pseudomonas sp. S9基因组中发现了大量ABCtansporter编码基因和细菌耐高浓度二价重金属离子Me2+的基因簇。这些基因的大量存在可能都与Pseudomonas sp. S9菌株适应红树林复杂生存环境相关。
整个Pseudomonas sp. S9基因组中存在7个与芳香族硫酸酯酶相关的蛋白。其中6个序列完整的芳香族硫酸酯酶都具有CX[PIA]XRX的保守结构域。4个Pseudomonas sp. S9的芳香族硫酸酯具有信号肽,表明这些芳香族硫酸酯不同于其它的Cry型芳香族硫酸酯酶,它们具有分泌到细胞外的潜力。首次报道了具有分泌功能的Cry型芳香族硫酸酯酶。
异源表达了菌株Pseudomonas sp. S9中的芳香族硫酸酯酶,对酶降解龙须菜硫酸根基团进行了相关功能研究。并且,通过对S9菌株全基因组文库的构建,获得了其中能够降解SDS的烷基型硫酸酯酶编码基因sdsAP,大肠杆菌中异源表达后的SdsAP具有很好的温度稳定性,扩展了烷基型硫酸酯酶的酶资源。
从海藻浒苔中筛选到一株高产琼脂酶的菌株LQ48,经鉴定LQ48为噬琼脂菌属淡黄色噬琼脂菌Agarivorans gilvus中的一个新菌株。克隆和重组表达了菌株Agarivorans sp. LQ48的GH16家族β-琼脂酶基因agaA,经过酶学性质分析,其具有广范围的pH稳定性。在琼脂酶AgaA研究基础上,通过LQ48菌株粗琼脂酶降解琼脂糖制备出不同聚合度的新琼寡糖混合物,利用分子筛成功地将新琼二糖NA2和新琼四糖NA4进行分离纯化。体外抗氧化性研究表明酶法制备的NA2和NA4均具有一定清除DPPH自由基、清除超氧阴离子自由基和清除羟基自由基的能力。
本研究的完成,为现行琼脂提取工艺的改进以及探索酶法生产高附加值琼脂产品的研究奠定理论基础。
The oceans are the Earth's largest ecosystem, covering 70% of our planet and providing goods and services for the majority of the world's population, particularly the algae. The extraction of agar is currently based on the procedure of alkali treatment, water washing, acid treatment, bleaching and water washing on raw algae materials, which not only cause serious environmental pollution, but also consumes vast quantities of water resources. On the other hand, the exported agar-relevant products are mainly low value-added agar, while processed agarose and agar oligosaccharides are imported with higher prices in China. Therefore, it is essential to improve the current process of extraction by applying the enzymatic treatment to degrade the sulfates of algae and produce the high value-added agar oligosaccharides, which will help to prevent environmental pollution and to gain the maximum economic benefits, thus promoting a faster and better development of the agar industry.
In this study, we isolated a sulfatase-producing bacterium S9 from the mangrove soil samples.16S rRNA analysis indicated that it belonged to genus Pseudomonas. The genome analysis suggests that the genome of Pseudomonas sp. S9 is consisted of a circular DNA. The draft genome excluding the gaps has a total of 4,796,832 bases comprising 4,665 predicted open reading frames (ORFs), with a G+C content of 56.52%. There are two rRNA operons and 56 predicted tRNA genes. In addition, the genome of Pseudomonas sp. S9 encodes amount of ABC tansporter genes and heavy metal tolerance genes. These are likely to bear systems to cope with toxic mangrove environment. According to the KEGG and COGs analysis, Pseudomonas sp. S9 is predicted that 38.3% function genes are related to the metabolism function. Moreover, it possesses the complete pathway of carbohydrate metabolism, lipid metabolism and amino acid metabolism. Additional, there are amounts of ABC transporter genes and heavy metal tolerance genes. These suggest Pseudomonas sp. S9 can cope with toxic metals in mangrove environment.
In S9 genome, seven predicted ORFs are annotated as arylsulfatases, six of whose sequences are complete. All these complete coding sequences have the CX[P/A]XRX motif, a characteristic of the cysteine-type sulfatases. Using SignalP to search for the possible signal peptides in cysteine-type sulfatases, we find four of them carrying a signal peptide which is accordant with that the arylsulfatases are exited in extracellular of Pseudomonas sp. S9. It is the first report of the extracellular cysteine-type sulfatases.
Meanwhile, arylsulfatase genes were cloned from strain S9 and expressed in E. coli BL21, and the recombinant enzyme was characterized to degrade the sulfate of polysaccharide from Gracilaria Lemaneiformis. That would provide a foundation for enzymatic degradation instead alkaline degradation in producing agar. Moreover, a novel alkylsulfatase gene, sdsAP, was cloned from a newly isolated bacterium Pseudomonas sp. S9 and expressed in Escherichia coli. The noticeable thermal stability makes SdsAP an ideal candidate for the application in disposing SDS-containing waste.
We described the isolation and identification of a new agarolytic bacterium Agarivorans sp. LQ48 from marine algae, and the cloning and expression of a novel endo-typeβ-agarase gene, agaA, in Escherichia coli. We also explored the characteristics of the purified recombinant agarase. On account of its noticeable pH stability, the enzyme can be utilized extensively in many kinds of industries. Then a series of neoagaroligosaccharides with different degree of polymerization were prepared by the crude agarases of strain LQ48. Neoagarobiose and neoagarotetraose were purified by Sephadex G-15 and identified by TLC. Furthermore, their antioxidant activity was investigated by various antioxidant assay in vitro systems, including DPPH radical scavenging, superoxide anion radical scavenging, hydrogen radical scavenging. The results indicated that neoagarobiose and neoagarotetraose producing by enzyme exhibited the antioxidant activities.
This study is the first step of improving the technique of agar production and exploring the methods of producing the neoagaroligosaccharides by agarases.
引文
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