CN114250211B - 一种甘露聚糖酶及其基因与应用 - Google Patents
一种甘露聚糖酶及其基因与应用 Download PDFInfo
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Abstract
本发明提供一种甘露聚糖酶,其氨基酸序列包括SEQ ID NO.1。本发明的甘露聚糖酶最适pH为6.0,在pH3.5‑7.0都有较高的酶活性;pH稳定性好;具有良好的抗蛋白酶的能力。其热稳定性高的特性,可使其在需求高温环境的工业生产上应用。本甘露聚糖酶可应用于饲料工业,降低或消除因粘度增加而引起的抗营养作用。此外,也可用作果汁饮料的澄清剂。还可与木聚糖酶协同作用破坏半纤维素与木质素之间的键结,减少漂白剂及氯的用量,进而保护环境,因而显示出巨大的潜力。
Description
技术领域
本发明属于基因工程及酶工程领域,具体涉及一种热稳定性提高的β-甘露聚糖酶突变体及其编码基因、包含该基因的重组载体和应用。
背景技术
β-甘露聚糖酶是一种能够降解β-1,4-甘露糖苷键的糖苷水解酶,在甘露聚糖降解酶系中发挥着重要作用,线状甘露聚糖、葡甘露聚糖、半乳甘露聚糖以及半乳葡甘露聚糖等甘露聚糖均可在其作用下水解为甘露糖或甘露寡糖。其底物甘露聚糖作为一种重要的植物半纤维素,是豆科植物细胞壁的主要组成部分,广泛存在于饲料原料中。对于体内不含甘露聚糖降解酶系的畜禽,饲料成分中的β-甘露聚糖会结合肠道内大量水分,使动物消化道内容物黏度升高,从而对胃肠的蠕动产生抵抗作用,同时还会减少破壁后养分与消化酶的接触,阻碍营养物质的消化吸收。而在饲料中添加β-甘露聚糖酶将甘露聚糖水解为低聚糖,能够有效的解决甘露聚糖带来的抗营养问题。另一方面,水解后的甘露低聚糖还是一种功能性营养物质,能够改善肠道菌群,提高饲料利用率,促进动物生长,改善动物的生产性能,从而进一步提高饲料的饲用价值,因此β-甘露聚糖酶在饲料产业中发挥重要的作用。
随着对β-甘露聚糖酶应用研究的深入开展,其在造纸、清洁、保健食品及生物技术等领域也得到了广泛的应用。在造纸领域,甘露聚糖酶可与木聚糖酶协同作用破坏半纤维素与木质素之间的键结,降低漂白剂及氯的用量;生活中常用到的发胶,护发素,调味酱等物品中普遍含有甘露聚糖用作增稠,采用含有耐碱甘露聚糖酶的洗涤剂可将其水解为易溶的小分子,使衣物等易漂洗。在食品领域,甘露聚糖酶能够用作果汁饮料的澄清剂等;此外甘露聚糖酶作为钻探过程中的优质生物破胶剂,具有成本低,破坏小,效率高等优点,因此在石油和天然气工业中也得到了应用。
基于甘露聚糖酶广阔的应用前景,甘露聚糖酶的研究逐渐成为国内外的研究热点。在饲料行业中,酶制剂生产后一般要经过高温造粒等工艺进行加工,成品常搭配饲料一同向动物饲喂,使得酶制剂需要先经由动物的胃部再到达肠道,才能发挥酶的作用。在此过程中,加工中的高温和动物肠道的消化酶都会对酶制剂产生不同程度的影响,降低其效果。因此开发一种热稳定性高,耐酸且对动物肠道消化酶有高耐受性的甘露聚糖酶具有重要的工业意义和经济价值。
发明内容
为了解决上述问题,本发明提供了一种中性甘露聚糖酶突变体,它具有耐高温,较宽的pH稳定性,对胰蛋白酶高耐受性等优点。
一方面,本发明提供了一种甘露聚糖酶。
所述的甘露聚糖酶为β-甘露聚糖酶。
所述的甘露聚糖酶氨基酸序列中包括SEQ ID NO.1。
所述的甘露聚糖酶N端包含信号肽序列,所述信号肽的序列为SEQ ID NO.2。
另一方面,本发明提供了一种基因。
所述的基因编码前述的甘露聚糖酶。
所述的基因上包括SEQ ID NO.3。
所述的基因上还包括编码信号肽的序列。
所述的信号肽为SEQ ID NO.2,所述的编码信号肽的序列包括根据密码子简并性衍生的任意可以编码SEQ ID NO.2的基因序列。
再一方面,本发明提供了一种重组载体。
所述的重组载体包含前述的基因。
优选地,所述的重组载体的基因***位点为EcoR I/Not I。
优选地,所述的重组载体为ppIC-man27s,即将本发明的甘露聚糖酶基因***到质粒pPIC9上的EcoR I和Not I限制性酶切位点之间,使该核苷酸序列位于AOX1启动子的下游并受其调控,得到重组酵母表达质粒pPIC9-man27s。
又一方面,本发明提供了一种基因工程细胞。
所述的基因工程细胞包含前述的重组载体。
优选地,所述的基因工程细胞的宿主细胞包括但不限于:毕赤酵母细胞、啤酒酵母细胞、多型逊酵母细胞、大肠杆菌、酵母菌、芽孢杆菌或乳酸杆菌,优选将为毕赤酵母细胞(Pichia pastoris)GS115。
又一方面,本发明提供了一种甘露聚糖酶的制备方法。
所述的制备方法中包括以下步骤:
1)用前述的重组载体转化宿主细胞,得重组菌株;
2)培养重组菌株,诱导重组甘露聚糖酶表达;
3)回收并纯化。
优选地,所述的步骤1)的宿主细胞包括但不限于:毕赤酵母细胞、啤酒酵母细胞、多型逊酵母细胞、大肠杆菌、酵母菌、芽孢杆菌或乳酸杆菌,优选将重组酵母表达质粒(即重组载体)转化毕赤酵母细胞(Pichia pastoris)GS115。
优选地,所述的步骤1)中的重组载体为pPIC-man27s。
优选地,所述的步骤1)和步骤2)中的重组菌株为GS115/man27s。
又一方面,本发明提供了前述的甘露聚糖酶和/或基因和/或重组载体和/或基因工程细胞在饲料、造纸、发胶、护发素、调味酱、保健食品、生物破胶剂、饮料澄清剂中的应用。
所述的在饲料中的应用为直接添加进饲料,或在饲料制备过程中进行酵解,或者其他可能的应用形式。
所述的在发胶或护发素中的应用为直接添加或者其他任意可行的形式的应用。
所述的在生物破胶剂中的应用为将甘露聚糖酶作为钻探过程中的生物破胶剂,或通过前述的基因载体或基因工程细胞制备甘露聚糖酶作为破胶剂;所述的应用为应用于石油或天然气工业或其他需要钻探的工业项目。
所述的在调味酱或保健食品或饮料澄清剂中的应用为直接添加或者其他任意可行的形式的应用。
所述的在造纸中的应用为与木聚糖酶协同作用破坏半纤维素与木质素之间的键结。
又一方面,本发明提供了一种酶制剂。
所述的酶制剂中包括前述的甘露聚糖酶和/或基因和/或重组载体和/或基因工程细胞。
又一方面,本发明提供了一种饲料。
所述的饲料中包括前述的甘露聚糖酶和/或基因和/或重组载体和/或基因工程细胞和/或酶制剂。
或者通过前述的甘露聚糖酶和/或基因和/或重组载体和/或基因工程细胞进行制备。
又一方面,本发明提供了一种食品或保健品添加剂。
所述的食品或保健品添加剂中包括前述的酶制剂,或通过前述的酶制剂进行制备。
又一方面,本发明提供了一种日用品添加剂。
所述的日用品添加剂包括前述的酶制剂,或通过前述的酶制剂进行制备。
本发明的有益效果:
本发明首先所要解决的技术问题是克服现有技术的不足,提供一种性质优良的、适合于在饲料、食品等行业中应用的耐热甘露聚糖酶。本发明的甘露聚糖酶最适pH为6.0,在pH3.5-7.0都有较高的酶活性(剩余酶活力>60%),可抗蛋白酶降解;pH稳定性好;具有良好的抗蛋白酶的能力。用胃蛋白酶和胰蛋白酶处理120分钟,酶活性分别提高了15%和10%。本发明的甘露聚糖酶热稳定性高,最适温度为60℃,在37℃仍具有50%左右的酶活力;可使其在需求高温环境的工业生产上应用。本甘露聚糖酶可应用于饲料工业,降低或消除因粘度增加而引起的抗营养作用。此外,甘露聚糖酶也可用作果汁饮料的澄清剂。还可与木聚糖酶协同作用破坏半纤维素与木质素之间的键结,减少漂白剂及氯的用量,进而保护环境,因而显示出巨大的潜力。
本发明的突变体Man27S,与Man27相比,该突变体的耐热性提高10℃,在75℃高温下保持5min仍维持80%酶活,而Man27在65℃下5min仅能维持60%的酶活。
附图说明
图1为重组甘露聚糖酶突变体的最适pH。
图2为重组甘露聚糖酶突变体的pH稳定性。
图3为重组甘露聚糖酶突变体的最适温度。
图4为重组甘露聚糖酶突变体的热稳定性。
具体实施方式
下面结合具体实施例,对本发明作进一步详细的阐述,下述实施例不用于限制本发明,仅用于说明本发明。以下实施例中所使用的实验方法如无特殊说明,实施例中未注明具体条件的实验方法,通常按照常规条件,下述实施例中所使用的材料、试剂等,如无特殊说明,均可从商业途径得到。
试验材料和试剂:
1、菌株及载体:本发明中甘露聚糖酶突变体基因man27s由北京睿博兴科生物技术有限公司合成获得,毕赤酵母表达载体pPIC9及菌株GS115购自于Invitrogen公司。
2、酶类及其它生化试剂:内切酶购自TaKaRa公司,连接酶购自Invitrogen公司。甘露聚糖购自Sigma公司,其它都为国产试剂(均可从普通生化试剂公司购买得到)。
3、培养基:
(1)酵母培养基YPD:1%蛋白胨、0.5%酵母提取物、1%葡萄糖、2%琼脂,pH7.0。
(2)大肠杆菌培养基LB:1%蛋白胨、0.5%酵母提取物、1%NaCl,pH7.0。
(3)BMGY培养基:1%酵母提取物、2%蛋白胨、1.34%YNB、0.00004%Biotin、1%甘油(V/V)。
(4)BMMY培养基:除以0.5%甲醇代替甘油,其余成份均与BMGY相同,pH4.0。
说明:以下实施例中未作具体说明的分子生物学实验方法,均参照《分子克隆实验指南》(第三版)J.萨姆布鲁克一书中所列的具体方法进行,或者按照试剂盒和产品说明书进行。
实施例1枯草芽孢杆菌Bacillus subtilis甘露聚糖酶突变体编码基因man27s的合成
本发明以来自于枯草芽孢杆菌(CGMCC NO.17882)的β-甘露聚糖酶Man27序列为SEQ ID NO.4,其包含信号肽的序列为SEQ ID NO.5,成熟的甘露聚糖酶Man27的理论分子量为38.3kDa;以编码SEQ ID NO.5的基因作为参考,对其序列进行如下突变:A18T,T83S,V136P,并在突变后序列的5’端与3’端分别加入EcoRI和NotI限制酶切位点,将序列发送至北京睿博兴科生物技术有限公司进行人工合成基因和基因载体。人工合成的甘露聚糖酶突变体氨基酸序列如SEQ ID NO.1所示,其核苷酸序列如SEQ ID NO.3所示,全长1014bp,命名该突变体为Man27S,基因命名为man27s。
实施例2甘露聚糖酶突变体基因man27s的克隆
合成好的基因载体以穿刺菌的形式保存,于超净台中用无菌牙签挑取穿刺菌,并置于含Amp(工作浓度:100μg/mL)抗生素的LB摇管中,37℃,220rpm过夜培养,次日按康为世纪质粒提取试剂盒PurePlasmid Mini Kit(CW0500)说明书步骤进行实验提取含有突变体基因的载体。
根据甘露聚糖酶突变体基因序列,设计并合成了如下引物:
P1:SEQ ID NO.6;
P2:SEQ ID NO.7;
以提取后的载体为模板进行PCR扩增。PCR反应参数为:94℃变性5min;然后94℃变性30sec,55℃退火30sec,72℃延伸1min,30个循环后72℃保温10min。得到一约1050bp片段,将该片段回收后与pMD19载体相连送北京睿博兴科生物技术有限公司测序,预测蛋白分子量为38.3kDa。
根据测序得到的核苷酸序列,通过DNAMan软件将得到的核苷酸序列与man27序列进行比对,确认A18T,T83S,V136P三处位置的突变无误。
实施例3重组甘露聚糖酶的制备
将表达载体pPIC9进行双酶切(EcoR I+Not I),同时将编码甘露聚糖酶突变体的基因man27s双酶切(EcoR I+Not I),酶切出编码成熟甘露聚糖酶的基因片段与表达载体pPIC9连接,获得含有甘露聚糖酶基因man27s的重组质粒pPIC-man27s并转化毕赤酵母GS115,获得重组毕赤酵母菌株GS115/man27s。
取含有重组质粒的GS115菌株与对照菌株(即未突变的菌株GS115/man27),接种于300mL BMGY培养液中,30℃250rpm振荡培养48h后,离心收集菌体。然后于150mL BMMY培养基重悬,30℃250rpm振荡培养。诱导72h后,离心收集上清,测定甘露聚糖酶的活力,即得重组甘露聚糖酶。
实施例4重组甘露聚糖酶Man27S的活性分析
DNS法检测重组甘露聚糖酶Man27S活性,设置未突变甘露聚糖酶Man27为对照,具体方法如下:
在pH5.5,37℃条件下,1mL的反应体系包括100μL适当的稀释酶液,900μL底物,反应10min,加入1.5mL DNS终止反应,沸水煮5min。冷却后540nm测定OD值。
甘露聚糖酶活性单位定义:在一定条件下,每分钟分解β-甘露聚糖生成lμmol还原糖所需的酶量为1个活性单位(U)。
重组甘露聚糖酶的表达量为4400U/mL,对照组甘露聚糖酶表达量为4370U/mL。SDS-PAGE结果表明,重组甘露聚糖酶在毕赤酵母中得到了表达。
实施例5重组甘露聚糖酶Man27S的性质测定
1、重组甘露聚糖酶Man27S的最适pH和pH稳定性的测定方法如下:
将对照组甘露聚糖酶Man27与重组表达的甘露聚糖酶Man27S在不同的pH条件下进行酶促反应分别测定最适pH。
所用缓冲液为pH0.5-2.2的KCl-HCl缓冲液,pH2.2-8.0的柠檬酸-磷酸氢二钠系列缓冲液及pH8.0-10.0的Tris-HCl系列缓冲液。
β-甘露聚糖酶Man27S在不同pH的缓冲体系,37℃下测定的pH适性结果(图1)表明,重组甘露聚糖酶Man27S的最适pH值为6.0,在pH3.5-7.0范围内,该酶能够维持60%以上的酶活力。
将酶液在不同pH值的缓冲液中37℃处理60min,测定酶活以研究酶的pH稳定性。结果(图2)表明重组甘露聚糖酶Man27S在pH3.0-10.0之间均很稳定,在此pH范围内处理60min仍能维持90%以上的酶活,说明此酶具有较好的pH稳定性,较对照组甘露聚糖酶Man27相比,重组甘露聚糖酶Man27S的pH稳定性有所提升。
2、甘露聚糖酶的最适温度及热稳定性测定方法如下:
甘露聚糖酶的最适温度的测定为在pH5.5条件下,分别测定不同温度(20-80℃)下的重组甘露聚糖酶Man27S与对照组甘露聚糖酶Man27的酶促反应活性,重组甘露聚糖酶Man27S的最适反应温度为60℃(图3)。
耐温性测定为甘露聚糖酶在不同温度下处理5min,再在60℃下进行酶活性测定。耐温性实验表明:重组甘露聚糖酶Man27S在75℃下处理5min,剩余酶活仍在80%以上(图4),而对照组甘露聚糖酶Man27在65℃处理5min即失去大部分酶活,表明重组甘露聚糖酶Man27S的热稳定性较对照组相比有明显提高。
3、重组甘露聚糖酶Man27S抗胃蛋白酶及胰蛋白酶能力测定如下:
用pH2.0的KCl-HCl缓冲液配制0.1mg/mL胃蛋白酶,pH7.0 Tris-HCl缓冲液配制0.1mg/mL胰蛋白酶。取pH2.0 KCl-HCl缓冲液稀释后的0.5mL纯化的酶液加入0.5mL胃蛋白酶,pH7.0 Tris-HCl缓冲液稀释后的0.5mL纯化的酶液加入0.5mL胰蛋白酶混合,蛋白酶/甘露聚糖酶(w/w)≈0.1,37℃保温60min和120min取样,在pH5.5及60℃条件下测定酶活性。
实验结果表明重组甘露聚糖酶Man27S用胃蛋白酶和胰蛋白酶处理120min后,胰蛋白酶处理后的Man27S的酶活比处理前提高了10%,胃蛋白酶处理后的Man27S甘露聚糖酶活性比处理前提高了15%,而对照组甘露聚糖酶经胰蛋白酶处理后酶活为处理前的95%,经胃蛋白酶处理后酶活为处理前的98%。以上结果表明突变后的重组甘露聚糖酶Man27S抗胃蛋白酶和胰蛋白酶的能力均有提高,具有良好的抗胃蛋白酶和胰蛋白酶的能力。
序列表
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<400> 5
Met Phe Lys Lys His Thr Ile Ser Leu Leu Ile Leu Phe Leu Leu Ala
1 5 10 15
Ser Ala Val Leu Ala Lys Pro Ile Glu Ala His Thr Val Ser Pro Val
20 25 30
Asn Pro Asn Ala Gln Gln Thr Thr Lys Ala Val Ile Asn Trp Leu Ala
35 40 45
His Leu Pro Asn Arg Thr Glu Asn Arg Val Leu Ser Gly Ala Phe Gly
50 55 60
Gly Tyr Ser His Asp Thr Phe Ser Met Ala Glu Ala Asp Arg Ile Arg
65 70 75 80
Ser Ala Thr Gly Gln Ser Pro Ala Ile Tyr Gly Cys Asp Tyr Ala Arg
85 90 95
Gly Trp Leu Glu Thr Ala Asn Ile Glu Asp Thr Ile Asp Val Ser Cys
100 105 110
Asn Ser Asn Leu Ile Ser Tyr Trp Lys Asn Gly Gly Ile Pro Gln Ile
115 120 125
Ser Leu His Leu Ala Asn Pro Ala Phe Gln Ser Gly His Phe Lys Thr
130 135 140
Pro Ile Thr Asn Asp Gln Tyr Lys Lys Ile Leu Asp Ser Ser Thr Val
145 150 155 160
Glu Gly Lys Arg Leu Asn Ala Met Leu Ser Lys Ile Ala Asp Gly Leu
165 170 175
Gln Glu Leu Glu Asn Gln Gly Val Pro Val Leu Phe Arg Pro Leu His
180 185 190
Glu Met Asn Gly Glu Trp Phe Trp Trp Gly Leu Thr Ser Tyr Asn Gln
195 200 205
Lys Asp Asn Glu Arg Ile Ser Leu Tyr Lys Gln Leu Tyr Lys Lys Ile
210 215 220
Tyr His Tyr Met Thr Asp Thr Arg Gly Leu Asp His Leu Ile Trp Val
225 230 235 240
Tyr Ser Pro Asp Ala Asn Arg Asp Phe Lys Thr Asp Phe Tyr Pro Gly
245 250 255
Ala Ser Tyr Val Asp Ile Val Gly Leu Asp Ala Tyr Phe Gln Asp Ala
260 265 270
Tyr Ser Ile Asn Gly Tyr Asp Gln Leu Thr Ala Leu Asn Lys Pro Phe
275 280 285
Ala Phe Thr Glu Val Gly Pro Gln Thr Ala Asn Gly Ser Phe Asp Tyr
290 295 300
Ser Leu Phe Ile Asn Ala Ile Lys Gln Arg Tyr Pro Lys Thr Ile Tyr
305 310 315 320
Phe Leu Ala Trp Asn Asp Glu Trp Ser Pro Ala Val Asn Lys Gly Ala
325 330 335
Ser Ala Leu Tyr His Asp Ser Trp Thr Leu Asn Lys Gly Glu Ile Trp
340 345 350
Asn Gly Asp Ser Leu Thr Pro Ile Val Glu
355 360
<210> 6
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
ccggaattca gatctcgctc 20
<210> 7
<211> 32
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
tttatagcgg ccgcttccac tatgggtgtt ag 32
Claims (13)
1.一种甘露聚糖酶,其特征在于,其氨基酸序列如SEQ ID NO.1所示。
2.根据权利要求1甘露聚糖酶,其特征在于,N端还包含信号肽序列,所述信号肽序列为SEQ ID NO.2。
3.一种基因,其特征在于,所述的基因编码权利要求1所述的甘露聚糖酶。
4.根据权利要求3所述的基因,其特征在于,所述的基因核苷酸序列如SEQ ID NO.3所示。
5.根据权利要求4所述的基因,其特征在于,所述的基因还包括编码信号肽的序列。
6.一种重组载体,其特征在于,包含权利要求3-5任一项所述的基因。
7.一种基因工程细胞,其特征在于,包含权利要求6所述的重组载体。
8.一种β-甘露聚糖酶的制备方法,其特征在于,包括以下步骤:
1)用权利要求6的重组载体转化宿主细胞,得重组菌株;
2)培养重组菌株,诱导重组甘露聚糖酶表达;
3)回收并纯化。
9.权利要求1-2任一项所述的甘露聚糖酶或权利要求3-5任一项所述的基因或权利要求6所述的重组载体或权利要求7所述的基因工程细胞在饲料、造纸、发胶、护发素、调味酱、保健食品、生物破胶剂、饮料澄清剂中的应用。
10.一种酶制剂,其特征在于,包括权利要求1-2任一项所述的甘露聚糖酶。
11.一种饲料,其特征在于,包括权利要求1-2任一项所述的甘露聚糖酶,或者通过权利要求1-2任一项所述的甘露聚糖酶进行制备。
12.一种食品添加剂,其特征在于,包括权利要求10所述的酶制剂,或通过权利要求10所述的酶制剂进行制备。
13.一种日用品添加剂,其特征在于,包括权利要求10所述的酶制剂,或通过权利要求10所述的酶制剂进行制备。
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CN101550422A (zh) * | 2009-05-15 | 2009-10-07 | 北京德宝群兴科技有限公司 | 一种β-甘露聚糖酶编码基因及其应用 |
CN105861466A (zh) * | 2016-05-15 | 2016-08-17 | 南京农业大学 | 通过基因工程改造获得的高活性甘露聚糖酶及其突变位点 |
CN110093331A (zh) * | 2019-05-06 | 2019-08-06 | 武汉轻工大学 | 一种高产的耐高温宽pH稳定性的甘露聚糖酶ManGold及基因与应用 |
CN111424048A (zh) * | 2020-06-09 | 2020-07-17 | 北京挑战农业科技有限公司 | 一种表达酸性β-甘露聚糖酶的基因及其载体和应用 |
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CN101550422A (zh) * | 2009-05-15 | 2009-10-07 | 北京德宝群兴科技有限公司 | 一种β-甘露聚糖酶编码基因及其应用 |
CN105861466A (zh) * | 2016-05-15 | 2016-08-17 | 南京农业大学 | 通过基因工程改造获得的高活性甘露聚糖酶及其突变位点 |
CN110093331A (zh) * | 2019-05-06 | 2019-08-06 | 武汉轻工大学 | 一种高产的耐高温宽pH稳定性的甘露聚糖酶ManGold及基因与应用 |
CN111424048A (zh) * | 2020-06-09 | 2020-07-17 | 北京挑战农业科技有限公司 | 一种表达酸性β-甘露聚糖酶的基因及其载体和应用 |
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