CN111454929A - 一种耐高温木聚糖酶基因及其应用 - Google Patents
一种耐高温木聚糖酶基因及其应用 Download PDFInfo
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- CN111454929A CN111454929A CN201910058604.4A CN201910058604A CN111454929A CN 111454929 A CN111454929 A CN 111454929A CN 201910058604 A CN201910058604 A CN 201910058604A CN 111454929 A CN111454929 A CN 111454929A
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- xylanase
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Abstract
本发明提供了一种耐高温木聚糖酶基因及其应用,该基因来自硫色曲酶,名为xyn10A,其序列如SEQ ID NO.1所示,该基因经偏好毕赤酵母的密码子优化后的核苷酸序列如SEQ ID NO.2所示,基于密码子优化后的xyn10A基因,本发明构建了高效分泌表达木聚糖酶xyn10A的毕赤酵母工程菌,该菌分泌表达的木聚糖酶具有热稳定性,最适催化温度70℃,最适pH值5.0,在pH4.0‑8.0之间保持高的相对活性,添加金属离子能提高木聚糖酶xyn10A的酶活,该酶作用于木聚糖30min的降解率为20.07%。本发明的木聚糖酶在造纸工业、饲料工业应用前景良好。
Description
技术领域
本发明涉及基因工程技术领域,具体地,涉及一种耐高温木聚糖酶基因及其应用。
背景技术
植植物细胞壁的组成成分包括纤维素、半纤维素以及木质素等,其中半纤维素的主要组成成分木聚糖,是一种丰富的生物质资源,是自然界除纤维素外最丰富的多糖。目前,降解木聚糖的方法有物理法、化学法和生物降解法。其中物理化学法各有其优缺点,但是酶解法相对于物理化学法,反应条件温和,反应专一。因此,物理法(热处理)以及化学法(酸碱处理)与酶解法结合被认为是降解木聚糖最有效的方法。木聚糖是一种多聚五碳糖,主链由β-D-1,4木糖苷键连接,侧链的组成较复杂,因此木聚糖的降解也需要一类酶的作用,即木聚糖酶,包括β-1,4-内切木聚糖酶,β-木糖苷酶、α-L-***糖苷酶、α-D-葡糖苷酸酶、乙酰基木聚糖酶和酚酸酯酶等。其中β-1,4-内切木聚糖酶作用于木聚糖的主链,是作用于木聚糖降解的关键酶。
在碳水化合物活性酶数据库(CAZy数据库)中,木聚糖酶基于催化结构序列的相似性具有10个不同的家族:GH5、GH8、GH10、GH11、GH16、GH26、GH30、GH43和GH62。木聚糖酶因其种类繁多,被广泛应用,可应用于工业上,如丹尼斯科美国公司2015年发现的一种GH10木聚糖酶可以用来提高油回收率;木聚糖酶也可用于制浆漂白中,可以水解木质纤维中的木聚糖,使植物的纤维结构松散从而有利于漂白剂的进入;另外,木聚糖酶也可应用于饲料中,木聚糖酶可降解植物原料中的木聚糖等抗营养因子,有利于动物机体对植物内营养物质的吸收。研究也表明,在禽类、猪、牛等反刍动物的饲料中添加木聚糖酶,可以提高营养物质的消化率。
研究表明,木聚糖酶的微生物来源主要有真菌和细菌,真菌产木聚糖酶主要有曲霉、青霉、木霉和酵母菌,而细菌产木聚糖酶的主要有大肠杆菌以及芽孢杆菌等。
发明内容
本发明的目的在于提供一种耐高温木聚糖酶基因及其应用。
本发明分析了硫色曲霉保藏编号CGMCC No.0608(该菌已在中国专利CN01141789.7中公开)。全基因组测序的结果,发现一条新的木聚糖酶基因序列xyn10A,基因序列1212bp,编码404个氨基酸,理论分子量43.64kDa。其中N末端22aa为信号肽序列。发明人利用NCBI网站BLAST在线比对分析,发现其与Aspergillus steynii IBT 23096的内切木聚糖酶(GenBank登录号:XP_024705365)的序列相似性最高为88%。本发明xyn10A基因编码的β-1,4-内切木聚糖酶属于糖苷水解酶(glycoside hydrolyse,GH)10家族。GH10家族的木聚糖酶具有较高的蛋白分子量、低等电点以及典型(α/β)8结构。
本发明提供一种木聚糖酶xyn10A,其氨基酸序列为:
a)SEQ ID No.3所示的氨基酸序列;或
b)SEQ ID No.3所示的氨基酸序列经替换、缺失和/或添加一个或几个氨基酸残基形成的具有同等功能的氨基酸序列。
本发明提供了编码所述木聚糖酶xyn10A的基因,是如下a)或b):
a)其核苷酸序列如序列表中SEQ ID No.1所示;或
b)由SEQ ID No.1所示核苷酸序列经取代一个或几个核苷酸,得到编码xyn10A的核苷酸序列。
优选地,本发明提供了经密码子优化后的对毕赤酵母偏好的木聚糖酶xyn10A的基因,其核苷酸序列如SEQ ID NO.2所示。
本发明进一步提供了含有上述基因的生物材料,所述生物材料为表达盒、载体、宿主细胞、或重组菌。
在本发明的实施例中,提供了含有SEQ ID NO.2所示基因的重组毕赤酵母菌,该毕赤酵母菌能够高效分泌表达木聚糖酶xyn10A。
含有所述生物材料的菌剂属于本发明的保护范围。
本发明提供了含有木聚糖酶xyn10A基因的生物材料,或含有本发明所述重组菌(毕赤酵母菌)的菌剂在制备耐高温木聚糖酶xyn10A中的应用。
本发明提供了含有所述木聚糖酶xyn10A的饲料。
本发明提供了所述的木聚糖酶xyn10A或其编码基因或含有其编码基因的生物材料在高温下降解木聚糖中的应用,所述高温为55-85℃,优选60-80℃。更优选70℃。在70℃条件下,
本发明提供了木聚糖酶xyn10A或其编码基因或含有该编码基因的生物材料在提高动物饲料中营养物质消化率中的应用,或在造纸工业中的应用。
本发明提供的耐高温木聚糖酶基因xyn10A来自硫色曲酶,该基因序列在现有技术中未见报道,其基因序列如SEQ ID NO.1所示,该基因经偏好毕赤酵母的密码子优化后的核苷酸序列如SEQ ID NO.2所示,基于该密码子优化后的xyn10A基因,本发明构建了高效分泌表达木聚糖酶xyn10A的毕赤酵母工程菌,该菌分泌表达的木聚糖酶具有热稳定性,最适催化温度70℃,相对活性达100%;最适pH值5.0,在pH 4.0-8.0之间稳定性较好,添加金属离子能提高木聚糖酶xyn10A的酶活,该酶作用于木聚糖30min的降解率为20.07%。本发明的木聚糖酶在造纸工业、饲料工业具有良好的应用前景。
附图说明
图1为木聚糖酶xyn10A的SDS-PAGE图,泳道A为对照,B为木聚糖酶xyn10A。
图2为木聚糖酶xyn10A的最适pH示意图。
图3为木聚糖酶xyn10A的最适温度示意图。
图4为木聚糖酶xyn10A的pH稳定性分析图。
图5为在50-90℃下的木聚糖酶xyn10A的温度稳定性分析图。
图6为木聚糖酶xyn10A的降解木聚糖的产物分析图。
具体实施方式
以下实施例用于进一步说明本发明的内容,但不应理解为对本发明的限制。在不背离本发明精神和实质的情况下,对本发明方法、步骤或条件所作的修改或替换,均属于本发明的范围。
若无特别指明,实施例中所用的技术手段为本领域技术人员所熟知的常规手段。
实施例1硫色曲霉的培养及其DNA的提取
将硫色曲霉CGMCC No.0608(已公开于中国专利CN01141789.7中)接种到灭菌的培养基(3.5g马铃薯葡萄糖琼脂培养基,5g D-麦芽糖,1g酵母提取物,2g蛋白胨,溶于100mL蒸馏水中,分装于50mL的三角瓶后再高压灭菌20min)中,30℃摇床培养72h。将菌体分离后置于液氮中研磨,利用真菌DNA提取试剂盒(Omega公司,美国)提取硫色曲霉全基因,并将硫色曲霉全基因进行基因组测序。
实施例2木聚糖酶xyn10A基因克隆
分析硫色曲霉全基因组测序的结果,发现一条新的木聚糖酶基因序列xyn10A,基因序列1212bp(如SEQ ID NO.1所示),编码404个氨基酸(如SEQ ID NO.3所示),理论分子量43.64kDa。其中N末端22aa为信号肽序列。利用NCBI网站BLAST在线比对分析,发现其与Aspergillus steynii IBT 23096的内切木聚糖酶(GenBank登录号:XP_024705365)的序列相似性最高为88%。
本发明对偏好毕赤酵母的xyn10A基因密码子优化后的核苷酸序列如SEQ ID NO.2所示。在北京擎科生物科技有限公司直接合成优化了的xyn10A序列,连接到pPICZαA载体上,转化入大肠杆菌(Escherichia coil,E.coil)TOP10感受态细胞中(北京天根生化科技有限公司),构建重组克隆质粒,通过含Zeocin(100μg/mL)(Invitrogen公司,美国)的LB固体培养基筛选菌落,并测序验证重组克隆质粒。
实施例3高效分泌表达木聚糖酶毕赤酵母工程菌株的构建
挑选转化好的大肠杆菌单菌落接种于LB液体培养基中,37℃摇床培养过夜,用质粒提取试剂盒(Omega公司,美国)提取高纯度重组克隆质粒,重组质粒用Sac I(TaKaRa公司,日本)线性化。
同时,制备毕赤酵母X-33感受态细胞。将线性化的表达制粒与毕赤酵母X-33感受态细胞混匀,转至0.2cm冰预冷的电转化杯中进行点击,点击完毕后,立即加入高压灭菌以及冰预冷的1mol/L等的山梨醇溶液,充分混匀,于28℃培养2-3h,将菌体悬液涂布含有Zeocin(100μg/mL)的YPDS固体平板培养基上,于28℃培养,直至长出清晰的菌落。挑取单菌落划线至含有Zeocin(100μg/mL)的YPD固定平板培养基上,28℃培养,所得菌株即为木聚糖酶xyn10A毕赤酵母表达工程菌株。
挑取重组毕赤酵母菌株,接种于含20mL BMGY(1%酵母提取物,2%蛋白胨,1.34%YNB(北京莱索宝科技有限公司),4×10-5%biotin,1%甘油,100mM pH 6.0磷酸盐缓冲液)培养基的250mL摇瓶中,于28℃、250rpm摇床培养至OD600值为2-6,5000rpm,4℃离心5min,弃上清,收集菌体,用BMMY(1%酵母提取物,2%蛋白胨,0.1mol/L磷酸缓冲液pH 6.0,1.34%YNB,4×10-5%biotin)重悬菌体,于28℃、250rpm摇床培养,再用终浓度为0.5%(V/V)的无水甲醇进行摇瓶诱导培养72h,每12h添加一次甲醇,离心取上清,4℃保存待测。
利用国标GBT23874-2009方法测木聚糖酶酶活,将木聚糖底物(Sigma公司,美国)溶液于70℃水浴预热5min,然后再分别吸取稀释适当比例酶液和水浴后木聚糖底物溶液各80ul充分混合,于70℃恒温水浴20min,加入200ul DNS终止反应,振荡混匀,沸水中煮5min,在流动的自然水下冷却至室温,用蒸馏水定容至1mL,利用酶标仪测定吸光度OD540,根据DNS标准曲线得到的OD540与还原性糖浓度的关系,计算得出木聚糖酶降解后还原性糖的浓度,从而计算得出木聚糖酶的酶活为100U/mL,实验过程中设置空白组,每个处理三个重复。酶活定义:在pH 5.0、70℃条件下,每分钟内底物降解释放1μmol还原性糖所需的酶量为一个酶活单位(U)。
实施例4木聚聚糖酶毕赤酵母工程菌株的酶学性质
将木聚糖酶毕赤酵母工程菌株摇瓶发酵后,离心过滤收集酶液。将收集的酶液经过SDS-PAGE分析,定性确定表达产物的大小在40kDa和55kDa之间,且接近于45kDa,与预计的木聚糖酶xyn10A分子量大小接近(如图1)。
将酶液适当稀释后,分别在55℃、60℃、65℃、70℃、75℃、80℃、85℃和90℃温度下测定并计算酶活,确定最适温度,实验均设置三个重复,下同。将酶液适当稀释后置于50℃下保温120min,前30min内每隔10min取一次样品,最后120min取一次样品;于60℃保温90min,每隔30min取一次样品,于70℃保温60min,前30min每隔10min取一次样品,之后每隔30min去一次样品;于80℃保温20min,每隔10min取一次样品,于90℃下保温10min,每隔5min取一次样品。每个温度设置一个对照组,保持温度为4℃,处理后于最适条件下测定酶活,确定酶液的温度稳定性。使用pH分别为2.5、3.0、3.5、4.0、4.5、5.0、5.5、6.0、6.5、7.0、7.5和8.0的柠檬酸-磷酸氢二钠缓冲液稀释酶液,底物也用相应pH的缓冲液配制成0.8%的底物溶液,在最适温度70℃条件下测定酶活,确定该酶的最适pH。将酶液分别置于上述pH的缓冲液中,室温处理30min后,最适条件下测定残余酶活,以最适pH的酶活作为对照组,确定该酶的pH稳定性。
实验结果表明,xyn10A的最适催化温度是70℃(图3),最适pH为5.0(图2);pH的稳定性较好,在pH 4-8之间保持较高的相对活性(图4),在50℃热处理120min,残余酶活为60%以上,60℃和70℃分别热处理30min,残余酶活均为40%以上,90℃处理5min,残余酶活在30%左右(图5)。
用pH 5的柠檬酸-磷酸氢二钠缓冲液配制成浓度分别为2mM、10mM和20mM的含有不同金属离子(见表1)和EDTA的化合物溶液,将酶液与金属离子以及EDTA等体积混合,使金属离子及EDTA终浓度为10mM,室温处理1h后,最适条件下测定酶活,以不添加金属离子和EDTA的酶液作为空白对照。
表1木聚糖酶xyn10A的不同浓度金属离子及EDTA耐受性下的相对酶活性/%
结果表明本发明的xyn10A酶对不同浓度金属离子以及EDTA常温作用1h的耐受性不一致,金属离子和EDTA终浓度为5mM时,所有的金属离子均提高了木聚糖酶xyn10A的酶活,而EDTA降低了木聚糖酶xyn10A的酶活。
金属离子和EDTA终浓度为10mM时,除Ca2+、Mg2+和EDTA外,所有的金属离子提高了木聚糖酶xyn10A的酶活,但提高酶活程度比5mM小(见表1)。
实施例5木聚糖酶xyn10A降解木聚糖的效果验证
用pH=5.0的柠檬酸-磷酸二氢钠缓冲液配制0.8%的木聚糖底物溶液,与适当比例稀释的实施例4获得的酶液分别反应0min、10min、20min和30min后,吸取一部分反应液,加入DNS终止反应,沸水中煮5min,在流动的自然水下冷却至室温,用蒸馏水定容,利用酶标仪测定OD540计算酶活。
另外一部分反应液直接放入4℃冰箱作为待测液。待测液稀释20倍后,用0.1μm滤膜过滤后进行液相离子色谱(DIONEX ICS-3000)分析,将标准品木二糖、木三糖、木四糖以及木五糖(上海甄淮生物科技有限公司,中国)稀释不同的浓度梯度,作为标准品计算产物木寡糖的含量,计算木聚糖酶的降解率。
由液相色谱图(图6)可以看出,xyn10A降解木聚糖的产物有木二糖、木三糖、木四糖和木五糖,根据标准品的液相色谱图,可以计算出木聚糖酶xyn10A作用于木聚糖30min的降解率为20.07%,随着降解时间由10min增加到30min,木二糖、木三糖、木四糖和木五糖的绝对含量在增加,木二糖的含量变化由降解10min时的37.19μg/mL,30min时增加到203.97μg/mL,占比由16.1%增加至26.76%;木三糖的绝对含量由41.05μg/mL增加到226.79μg/mL,占比由26.04%增加到31.58%;木四糖的绝对含量由41.90μg/mL增加到172.78μg/mL,占比由26.58%降低到24.06%;木五糖的绝对含量由49.32μg/mL增加到126.46μg/mL,占比由31.28%降低至17.61%。
液相色谱参数:色谱柱:00H-0138-K0 Column(250mm×4mm);保护柱:CarboPacPA10(50mm×4mm),淋洗液A为250mmol/L的NaOH,淋洗液B为超纯水,液体流速:1mL/min;进样体积:100μL。
虽然,上文中已经用一般性说明、具体实施方式及试验,对本发明作了详尽的描述,但在本发明基础上,可以对之作一些修改或改进,这对本领域技术人员而言是显而易见的。因此,在不偏离本发明精神的基础上所做的这些修改或改进,均属于本发明要求保护的范围。
序列表
<110> 中国农业大学
<120> 一种耐高温木聚糖酶基因及其应用
<130> KHP191110383.7
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ggcggcaaca aggccgctgg tgcggtcagg atcgtcaagc tgatccagtc gtacggcgtg 660
aagatcgacg gagtcggtct ccagggccac ttcactgtcg gtaacatccc tggtaagaat 720
gacctggcca gtaccttgaa aacctacacc gtgctcggtg tcgaggtggc gtacaccgaa 780
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ttcggtcaaa ttactccagg taactctcaa aagtgggatt ctactgaacc atctcaaaac 180
accttctctt tcactaaggg tgacgctatt gctgatttgg ctaaggctaa caaccaagtt 240
ttgcgttgcc ataacttggt ttggcatcaa caattgccaa actgggttac ttctggttct 300
tggactaacg ctactttgac tgctgttttg aagaaccaca ttaccaacgt cgttaagcac 360
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ggtggtaaca aggctgctgg tgctgttaga atcgtcaagt tgatccaatc ttacggcgtc 600
aagattgatg gtgttggttt gcaaggtcat tttaccgttg gtaacatccc aggtaagaac 660
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tcccaatgct tgtagtctag a 1161
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Glu Phe Ala Gly Leu His Gln Ala Ala Val Ala Lys Gly Leu Asn Tyr
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Phe Gly Thr Ala Thr Asp Asn Pro Glu Leu Thr Asp Ile Pro Tyr Val
20 25 30
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100 105 110
His Ile Thr Asn Val Val Lys His Tyr Lys Gly Arg Cys Tyr Ala Trp
115 120 125
Asp Val Val Asn Glu Ala Leu Asn Glu Asp Gly Ser Tyr Arg Asp Ser
130 135 140
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145 150 155 160
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165 170 175
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180 185 190
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195 200 205
Gly His Phe Thr Val Gly Asn Ile Pro Gly Lys Asn Asp Leu Ala Ser
210 215 220
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225 230 235 240
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290 295 300
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305 310 315 320
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325 330 335
Ser Thr Val Ser Ala Pro His Thr Thr Ser Thr Asn Val Ala Gln Lys
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Ser Gly Thr Thr Cys Thr Lys Leu Asn Asp Trp Tyr Ser Gln Cys Leu
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1 5 10 15
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20 25 30
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35 40 45
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50 55 60
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65 70 75 80
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85 90 95
Gly Ser Trp Thr Asn Ala Thr Leu Thr Ala Val Leu Lys Asn His Ile
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Thr Asn Val Val Lys His Tyr Lys Gly Arg Cys Tyr Ala Trp Asp Val
115 120 125
Val Asn Glu Ala Leu Asn Glu Asp Gly Ser Tyr Arg Asp Ser Ile Phe
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145 150 155 160
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180 185 190
Ile Gln Ser Tyr Gly Val Lys Ile Asp Gly Val Gly Leu Gln Gly His
195 200 205
Phe Thr Val Gly Asn Ile Pro Gly Lys Asn Asp Leu Ala Ser Thr Leu
210 215 220
Lys Thr Tyr Thr Val Leu Gly Val Glu Val Ala Tyr Thr Glu Val Asp
225 230 235 240
Val Arg Met Glu Thr Pro Ala Thr Asp Ala Lys Leu Ala Gln Gln Ser
245 250 255
Ile Asp Tyr Gln Asn Leu Val Gln Ala Cys Val Glu Thr Pro Lys Cys
260 265 270
Val Gly Phe Thr Ile Trp Asp Trp Thr Asp Lys Tyr Ser Trp Val Pro
275 280 285
Ser Thr Phe Pro Gly Gln Gly Ala Ala Cys Pro Trp Asp Glu Asn Leu
290 295 300
Lys Lys Lys Pro Ala Tyr Thr Gly Leu Leu Lys Gly Leu Gly Gly Asn
305 310 315 320
Arg Ser Glu Ser Ser Ser Ser Ser Ser Ser Ser Thr Pro Thr Ser Thr
325 330 335
Val Ser Ala Pro His Thr Thr Ser Thr Asn Val Ala Gln Lys Trp Gly
340 345 350
Gln Cys Gly Gly Asn Asn Trp Thr Gly Pro Thr Thr Cys Val Ser Gly
355 360 365
Thr Thr Cys Thr Lys Leu Asn Asp Trp Tyr Ser Gln Cys Leu
370 375 380
Claims (10)
1.一种木聚糖酶xyn10A,其氨基酸序列为:
a)SEQ ID No.3所示的氨基酸序列;或
b)SEQ ID No.3所示的氨基酸序列经替换、缺失和/或添加一个或几个氨基酸残基形成的具有同等功能的氨基酸序列。
2.编码权利要求1所述木聚糖酶xyn10A的基因,其特征在于,是如下a)或b):
a)其核苷酸序列如序列表中SEQ ID No.1所示;或
b)由SEQ ID No.1所示核苷酸序列经取代一个或几个核苷酸,得到编码xyn10A的核苷酸序列。
3.如权利要求2所述的基因,其特征在于,其核苷酸序列如SEQ ID NO.2所示。
4.含有权利要求2或3所述基因的生物材料,所述生物材料为表达盒、载体、宿主细胞、或重组菌。
5.如权利要求4所述的生物材料,其为重组毕赤酵母菌。
6.含有权利要求5所述生物材料的菌剂。
7.权利要求4或5所述的生物材料,或权利要求6所述的菌剂在制备耐高温木聚糖酶xyn10A中的应用。
8.含有权利要求1所述木聚糖酶xyn10A的饲料。
9.权利要求1所述的木聚糖酶xyn10A或权利要求2或3所述的编码基因或权利要求4或5所述的生物材料在高温下降解木聚糖中的应用,所述高温为55-85℃,优选60-80℃。
10.权利要求1所述的木聚糖酶xyn10A或权利要求2或3所述的编码基因或权利要求4或5所述的生物材料在提高动物饲料中营养物质消化率中的应用,或在造纸工业中的应用。
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