CN107267529A - 一种锌指蛋白转录因子基因RkMSN4及其应用 - Google Patents
一种锌指蛋白转录因子基因RkMSN4及其应用 Download PDFInfo
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Abstract
本发明公开了一种锌指蛋白转录因子基因RkMSN4，该基因的核苷酸序列如SEQ ID NO：1所示，该基因编码的氨基酸序列如SEQ ID NO：2所示；该基因是从红冬孢酵母YM25235中分离得到调控红冬孢酵母中催化油酸转化为亚油酸（LA）和α‑亚麻酸（ALA）的Δ12/15‑脂肪酸脱氢酶基因RKD12的转录因子；为研究红冬孢酵母抗低温环境机理提供参考，同时通过基因工程手段对微生物进行改造来提高多不饱和脂肪酸亚油酸和α‑亚麻酸的含量，对PUFAs的工业化生产提供良好的应用前景和经济效益，为大规模商业化生产PUFAs奠定基础。
Description
技术领域
本发明属于生物技术领域和遗传工程领域，涉及一种锌指蛋白转录因子基因RKMSN4及其重组表达载体，具体涉及从红冬孢酵母（Rhodosporidium kratochvilovae）YM25235中克隆的锌指蛋白转录因子基因RkMSN4及将该基因连接到载体转入到酵母菌中，并对其调节酵母菌生产多不饱和脂肪酸的能力进行研究，为多不饱和脂肪酸的大规模生产提供参考。
背景技术
多不饱和脂肪酸（polyunsaturated fatty acids，PUFAs）是指含有两个或两个以上 C = C 且碳链长度为 18～22 个碳原子的脂肪酸。根据第一个 C = C出现在从甲基原子数起第 3个或第 6个碳原子上，通常分为 n-3 和 n-6 系列。多不饱和脂肪酸具有多种重要的生理功能，包括可降低冠心病、神经退行性疾病的风险。此外，PUFAs 还可以作为很多具有生物活性物质的前体物质，如***素、白三烯和血栓素，这些生物活性物质可以有效 缓解炎症、疼痛和发烧。
长期以来人们都在研究如何从动植物油脂中提取PUFAs，但因为动植物的生长随着季节、地理位置等的影响而不断发生变化，所以PUFAs含量和构成也随之而变，况且从动植物中提取PUFAs的成本高，周期长，不能适应市场的需要。另外动植物油脂资源含油量及不饱和脂肪酸类型、比例均受到一定的限制。因此，近年来人们一直在探索利用PUFAs的新来源---即利用微生物技术来生产PUFAs。微生物具有发酵周期短、油脂含量高、培养成本低、不受原料控制、对环境的适应性强、生物转化率高等优点。近年来对于脂质积累的研究主要集中于单细胞微生物，如酵母和微藻。
锌指蛋白MSN4是一类转录因子，很多研究表明，不同环境胁迫条件下其能促进宿主细胞中胁迫适应相关基因的转录，从而促进宿主对高盐、高渗透压等环境胁迫适应性，但是目前没有锌指蛋白MSN4低温条件下促进多不饱和脂肪酸合成的研究报道。
发明内容
本发明的目的是提供一种锌指蛋白转录因子基因RkMSN4，其参与调节低温环境下红冬孢酵母YM25235抗低温环境关键酶RkD12脂肪酸脱氢酶的活性，使酵母大量产生多不饱和脂肪酸抵御低温环境。该基因是从红冬孢酵母（Rhodosporidium kratochvilovae）YM25235中分离得到，该基因核苷酸序列如SEQ ID NO：1所示，该基因序列长为2625bp（碱基），该基因编码的氨基酸序列如SEQ ID NO：2所示的多肽或其片段，共编码874个氨基酸。
本发明另一目的是提供一种含有锌指转录因子RkMSN4的重组表达载体，是将SEQID NO：1所示基因直接与原核表达载体pRH2034连接所构建的重组载体pRH2034RkMSN4。
本发明另一目的是将锌指蛋白转录因子基因RkMSN4应用在低温下生产多不饱和脂肪酸中。
本发明从红冬孢酵母（Rhodosporidium kratochvilovae）YM25235的总RNA基因中分离得到调控红冬孢酵母抗低温环境锌指蛋白转录因子基因RkMSN4，该基因全长2625bp。研究表明锌指转录因子RkMSN4在低温下能提高红冬孢酵母YM25235中催化油酸转化为亚油酸和亚麻酸的Δ12/15-脂肪酸脱氢酶基因RKD12基因的mRNA转录水平的提高，从而引起细胞脂中PUFAs的合成量显著增加；本研究结果有助于阐明产油酵母-红冬孢酵母YM25235中PUFAs低温合成调节机制，为揭示微生物低温适应性的机制提供参考，将有助于通过基因工程手段对其进行改造来提高多不饱和脂肪酸-亚油酸（LA）和α-亚麻酸（ALA）的含量，对多不饱和脂肪酸的工业化生产提供有良好的应用前景和经济效益，为大规模商业化生产多不饱和脂肪酸奠定基础。
附图说明
图1为本发明的红冬孢酵母YM25235 锌指蛋白转录因子RkMSN4基因PCR扩增图；
图2为利用本发明的红冬孢酵母YM25235 锌指蛋白转录因子RkMSN4基因构建的红冬孢酵母表达载体pRH2034RkMSN4；
图3为本发明所构建的重组表达质粒pRH2034RkMSN4的酶切分析图；其中：1为DNAMaker；2为空质粒pRH2034 BamH I、EcoR V双酶切对照；3、4为重组质粒pRH2034RkMSN4BamH I、EcoR V双酶切；5为RkMSN4基因PCR产物；
图4 为低温条件下 RkMSN4基因转化的红冬孢酵母YM25235细胞脂肪酸气相色谱分析；A: 30℃培养的YM25235/pRH2304；B: 30℃培养的YM25235/ pRH2034RkMSN4；
图5为低温条件下 RkMSN4基因转化的红冬孢酵母YM25235细胞脂肪酸气相色谱分析；A: 15℃培养的YM25235/pRH2304；B: 15℃培养的YM25235/ pRH2034RkMSN4。
具体实施方式
下面结合附图和实施例对本发明作进一步详细说明，但本发明保护范围不局限于所述内容，实施例中使用的试剂和方法，如无特殊说明，均采用常规试剂和使用常规方法。
实施例1：红冬孢酵母（Rhodosporidium kratochvilovae）YM25235锌指蛋白转录因子RkMSN4基因克隆
采用OMEGA试剂盒E.Z.N.A Fungal RNA Kit提取红冬孢酵母YM25235总RNA，反转录试剂盒Takara First Strand cDNA Synthesis Kit合成cDNA。根据红冬孢酵母YM25235的转录组序列设计特异性引物（引物1和引物2）进行PCR扩增；反应所用引物、组份和扩增条件如下：
引物1：RkMSN4-F：5’-AATGGATCCATGGCGCCTGCTCCCCGT-3’（SEQ ID NO:3）
引物2：RkMSN4-R：5’-ATCGATATCTCACGGCATCGCCCACACCT-3’（SEQ ID NO:4）
（GGATCC为BamH I酶切位点，GATATC为EcoRV酶切位点）
PCR扩增体系如下（50 μL）：
PCR扩增条件为：95℃预变性5min，95℃ 变性30s、 64℃退火 30s、72℃ 延伸3min进行30个循环，最后72℃延伸10min。反应完后取产物1μL，然后在浓度为1%的琼脂糖凝胶中进行电泳分析。结果如图1所示，扩增获得大约2700bp大小的片段，然后用百泰克生物技术有限公司多动能DNA纯化回收试剂盒回收目的片段，然后将PCR扩增得到的目的基因连接到pMD18-T上，连接产物转化至大肠杆菌DH5α感受态细胞中，用含有氨苄青霉素（Amp+）的LB固体平板进行筛选，挑取平板上的转化子进行菌落PCR筛选阳性克隆，然后送去上海生工测序。测序结果表明，获得一段2625bp长的序列，编码874个氨基酸，命名为RkMSN4，序列组成如SEQ ID NO:1所示的核苷酸序列和氨基酸序列。
实施例2：重组表达质粒pRH2034RkMSN4的构建
采用实施例1中测cDNA为模板进行PCR扩增，反应所以引物组合、反应组分和扩增条件如下：
引物1：RkMSN4-F：5’-AATGGATCCATGGCGCCTGCTCCCCGT-3’（SEQ ID NO:3）
引物2：RkMSN4-R：5’-ATCGATATCTCACGGCATCGCCCACACCT-3’（SEQ ID NO:4）
（GGATCC为BamH I酶切位点，GATATC为EcoRV酶切位点）
PCR扩增体系如下（50 μL）：
PCR扩增条件为：95℃预变性5min，95℃ 变性30s、 64℃退火 30s、72℃ 延伸3min进行30个循环，最后72℃延伸10min。将上述PCR获得的基因片段和表达载体pRH2034用BamH I和 EcoRV核酸内切酶进行双酶切，用百泰克生物技术有限公司多动能DNA纯化回收试剂盒回收酶切片段，并用T4连接酶在16℃连接过夜，将片段连接到表达载体 pRH2304上；连接产物转化大肠杆菌DH5α，用含有壮观霉素（Spectinomycin）的LB固体平板进行筛选，挑取平板上的转化子进行菌落PCR，筛选阳性克隆，同时进一步对转化子进行双酶切验证分析，结果显示，如图3第3、4泳道所示，用EcoRV和BamH I双酶切，重组质粒产生两条带，小分子条带与泳道5中该基因的PCR产物大小一致，大分子条带与泳道2中用相同酶切质粒pRH2034后产生的条带大小一致，这表明所构建的重组质粒正确，进一步测序分析也证明这一点；将重组表达质粒命名为pRH2034RkMSN4，该质粒图谱如图2所示。
实施例3：RkMSN4基因与低温环境中合成多不饱和脂肪酸实验
1、农杆菌介导转化红冬孢酵母YM25235
采用农杆菌介导转化法将重组质粒pRH2034RkMSN4转化至红冬孢酵母YM25235菌株，以含潮霉素B（HygromycinB）终浓度为150 µg/mL的 YPD 培养基筛选转化子；提取基因组DNAPCR验证转化子。
2、RKMSN4基因与红冬孢酵母脂肪酸含量变化的分析
将转基因菌株YM25235/ pRH2034RkMSN4及对照菌株YM25235/pRH2304分别先在30℃培养24 h后，一份30℃继续培养24 h，另一份立即转移至15℃冷处理培养24 h；分别提取冷处理后4株菌的细胞中总脂肪酸，并进行甲酯化。将样品进行气相色谱分析，利用面积归一法计算菌株脂肪酸的含量；4株菌的脂肪酸气相色谱分析图谱如图4、5所示，在图中保留时间约为11 min为饱和脂肪酸C18:1 油酸OA，保留时间约为12.5 min为不饱和脂肪酸C18:2亚油酸LA，保留时间约为15 min为不饱和脂肪酸C18:3亚麻酸ALA。从图4、5，我们可以发现，当转基因菌株YM25235/pRH2034RkMSN4和对照菌株YM25235/pRH2304在30 ℃条件下，油酸（OA）、LA及ALA的峰面积大小基本一致；而当转基因菌株YM25235/ pRH2034RkMSN4和对照菌株YM25235/pRH2304在15℃培养后，油酸（OA）的峰面积大小均呈下降趋势，LA及ALA的峰面积大小均呈上升趋势，且转基因菌株YM25235/ pRH2034RkMSN4的LA及ALA的峰面积均大于对照菌株YM25235/pRH2304 ALA的峰面积（图4A: 30℃培养的YM25235/pRH2304，B: 30℃培养的YM25235/ pRH2034RkMSN4；图5A: 15℃培养的YM25235/pRH2304，B: 15℃培养的YM25235/ pRH2034RkMSN4）。
根据下表中脂肪酸含量数据，我们可以计算得出，在经15℃培养后，转基因菌株YM25235/ pRH2034RkMSN4中LA和ALA含量分别为31.56%和17.32%，相对于同一菌株30℃培养条件下LA和ALA的含量分别为16.81 %和6.14%，变化不明显，但是相比较于15℃培养条件下对照菌株YM25235/pRH2304中LA和ALA的含量（分别为24.18%和8.73%）分别提高了1.31倍和1.98倍，而且OA的含量减少了0.58倍。这些结果表明，在红冬孢酵母YM25235菌株中过表达RkMSN4基因会引起低温条件下LA及ALA两种多不饱和脂肪酸含量的增加。
表1：15℃条件下RkMSN4转基因红冬孢酵母YM25235细胞总脂肪酸含量
。
序列表
<110> 昆明理工大学
<120> 一种锌指蛋白转录因子基因RkMSN4及其应用
<160> 4
<170> PatentIn version 3.3
<210> 1
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<213> Rhodosporidium kratochvilovae
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atggcgcctg ctccccgtgg ttctgccagc gggcagacct tcccctgccc cgactgcgac 60
aaggtcttct cccgcaagga gtacatggcg cgacattacc ggtcgaggca tagcaaggag 120
aagcccttcc aatgcgagta ctgcgaacac gccttcagcc gcagtgacct gctgaggcgt 180
caccacaaaa cgtgcagcca ggctaaggcg gcgaggggcg agggcgactc gccggatctt 240
ccgcgcgagt cgcccaccga gcctgccgcg cccgtagcgt cgacctccgc catgcccgtc 300
gacatactgt acccgccgta cccaccgccc gtcgcctcgt cctcgtccgc cgttcctgcg 360
ccgacctact cgccatatga cacaggcgcg caggacctgc ctgtcccccc agccagcttc 420
ctcgacctcg tctcttcgca accgtaccct gcgccctccc cgtccctccc cgccttctcg 480
cccaacatgg cgagcctgta tcccatccca ttgaggaccg gcagcacagc ctcgcccgcc 540
tcgagcggca tgtccggctc gccgttctcg cgcgccacgt acgcctcgac gaccggcaca 600
tcgcccgaca tggcgatgca gaaggagctg ccgctcacgc tccctgacgt gatggcttcc 660
atctcggcga cgacgagcgc gccgcgccct caaccgccgc aggagcagca agcgatgccg 720
acgctcgccg cgctcgaccc gtacggcgcc gcgccgcagc cttcgtccgc attcaaggcc 780
ggcgtcacgc agcacgatac gctcgcgccc ggcctgtcgc gcacgggcag cttcaccaag 840
gacgaagtgc tcgcgtccga ggtcctgcag gacctgatgc gcacgccgtt cggcgcgccg 900
taccccgcgc cgccagcgag cacgagcggc gtgccctggc acggcgcgaa ggcggcgcag 960
cgcgcggcgg agcagcccga gtcggcgcag gcgaacgaag ctgtcagcct ggtcgactcg 1020
tccggcggcg actgggcttt cagcctcggc ggcggcgcgg ttctgcccat cggcgggttc 1080
ggcgggcccg tctcaaacaa gctggaggag acgccagcgg cgcagcagct cgccgagtac 1140
ttcaataagg gcggcgtcgg cggcatcacg gcgctcgacc tcggcttcac ggtcgagccg 1200
agcctgtggc ctgagtggat cacgatggag ccgaagccgg tgcacgacga catgaagcgg 1260
tggtggctgc cagagcagaa gttctgcctc ggctatctct acccttggca tgtcccgccc 1320
cttcccgttc tctccggata cgcccgcaag gctacggagc agctcctcgc agctgtgccc 1380
gtcctgcatg gtccgtccgc ggtcatgacc gagctaccga cgcacacggc gtttgcgctg 1440
accgtcgccg gcggggcgta cgagcccgag ggccagagct tcagtaacga gatgctcgtc 1500
gagaagaggg tgttcctcgt gcgcggcttc caggagaagg acaagagctg ggaggaccgc 1560
ttcgcgtcac tccagtcgct cctgctctac cagcttctcg gcatcttcca ccgcgacgag 1620
cagcagcgcc tgctctccca ctccttccat tcggcgctcg tctacatgtt tcgcgcgctc 1680
gacctcccga accgcgtaac gcagaccgcg ctcgtcaagc cgcgggcaga catgcatggt 1740
ggggagctcg agaaggcgtg gaaggagtgg atcgaggtcg aaacccggcg gcgagtcgcg 1800
ttcatcgttt tcctcatcga cctcgagcac gcggcggcga ccgagacgcc gcagctcctc 1860
gcgctcagcg acctcgacct cgaccttccc gcctccgagc gcgcgtggaa ggcagagaac 1920
gccgccgaat ggctcgatcg cagtacctcg cccctctacc ccgaaaccat ctccttcctc 1980
gccgccatcc gcgccctcct ctcgactacc ccgcccgagc ccttctccgc cgcaggcgtt 2040
ctcctcgccg aactcggccg cctctcgtcc ttcccgctcc tcatcctctc gcggacgctc 2100
tcgtacctcg agcgcaagac gcaggaggcg ctcgcgcaaa tcgacccgtt caagagcctc 2160
ctcggcggcc tcggcgtcct tgagggccgc gagaccgaga atcgcgccgt cctcgagcgc 2220
attcgtcgcg gccgcgaagt cctgcgccgc ctgcctggcg ggattgcgcg cgggggtgga 2280
gagggctggt tcaacgagat catcccctcg gccaaggatt tcgtcccagc gtcggactcg 2340
cccgcagcgt cgagttcgag cgcgggcacc tccccgcggg acgcgttctc ctcctcctcc 2400
gcaagcggga cggcatcgac atttccctcg ccgcaagacg cgcccgcctc gctcgaggag 2460
ctcttcgccg agttcgacga gcagccgtac cgcccgttcc atggcgcggg cggcgcgacg 2520
cccggcgaga cgtacgagca ggcgcaggag cgcctgcgca agcatgccga gcggcgcgtg 2580
cgcgatgcgc agtccgcact gcccgaggtg tgggcgatgc cgtga 2625
<210> 2
<211> 874
<212> PRT
<213> Rhodosporidium kratochvilovae
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MET Ala Pro Ala Pro Arg Gly Ser Ala Ser Gly Gln Thr Phe Pro
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Cys Pro Asp Cys Asp Lys Val Phe Ser Arg Lys Glu Tyr MET Ala
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Glu Tyr Cys Glu His Ala Phe Ser Arg Ser Asp Leu Leu Arg Arg
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His His Lys Thr Cys Ser Gln Ala Lys Ala Ala Arg Gly Glu Gly
70
Asp Ser Pro Asp Leu Pro Arg Glu Ser Pro Thr Glu Pro Ala Ala
80 90
Pro Val Ala Ser Thr Ser Ala MET Pro Val Asp Ile Leu Tyr Pro
100
Pro Tyr Pro Pro Pro Val Ala Ser Ser Ser Ser Ala Val Pro Ala
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Pro Thr Tyr Ser Pro Tyr Asp Thr Gly Ala Gln Asp Leu Pro Val
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Pro Pro Ala Ser Phe Leu Asp Leu Val Ser Ser Gln Pro Tyr Pro
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Ala Pro Ser Pro Ser Leu Pro Ala Phe Ser Pro Asn MET Ala Ser
160
Leu Tyr Pro Ile Pro Leu Arg Thr Gly Ser Thr Ala Ser Pro Ala
170 180
Ser Ser Gly MET Ser Gly Ser Pro Phe Ser Arg Ala Thr Tyr Ala
190
Ser Thr Thr Gly Thr Ser Pro Asp MET Ala MET Gln Lys Glu Leu
200
Pro Leu Thr Leu Pro Asp Val MET Ala Ser Ile Ser Ala Thr Thr
220
Ser Ala Pro Arg Pro Gln Pro Pro Gln Glu Gln Gln Ala MET Pro
230
Thr Leu Ala Ala Leu Asp Pro Tyr Gly Ala Ala Pro Gln Pro Ser
250
Ser Ala Phe Lys Ala Gly Val Thr Gln His Asp Thr Leu Ala Pro
260
Gly Leu Ser Arg Thr Gly Ser Phe Thr Lys Asp Glu Val Leu Ala
280
Ser Glu Val Leu Gln Asp Leu MET Arg Thr Pro Phe Gly Ala Pro
290
Tyr Pro Ala Pro Pro Ala Ser Thr Ser Gly Val Pro Trp His Gly
310
Ala Lys Ala Ala Gln Arg Ala Ala Glu Gln Pro Glu Ser Ala Gln
320
Ala Asn Glu Ala Val Ser Leu Val Asp Ser Ser Gly Gly Asp Trp
340
Ala Phe Ser Leu Gly Gly Gly Ala Val Leu Pro Ile Gly Gly Phe
350
Gly Gly Pro Val Ser Asn Lys Leu Glu Glu Thr Pro Ala Ala Gln
370
Gln Leu Ala Glu Tyr Phe Asn Lys Gly Gly Val Gly Gly Ile Thr
380
Ala Leu Asp Leu Gly Phe Thr Val Glu Pro Ser Leu Trp Pro Glu
400
Trp Ile Thr MET Glu Pro Lys Pro Val His Asp Asp MET Lys Arg
410
Trp Trp Leu Pro Glu Gln Lys Phe Cys Leu Gly Tyr Leu Tyr Pro
430
Trp His Val Pro Pro Leu Pro Val Leu Ser Gly Tyr Ala Arg Lys
440
Ala Thr Glu Gln Leu Leu Ala Ala Val Pro Val Leu His Gly Pro
460
Ser Ala Val MET Thr Glu Leu Pro Thr His Thr Ala Phe Ala Leu
470
Thr Val Ala Gly Gly Ala Tyr Glu Pro Glu Gly Gln Ser Phe Ser
490
Asn Glu MET Leu Val Glu Lys Arg Val Phe Leu Val Arg Gly Phe
500
Gln Glu Lys Asp Lys Ser Trp Glu Asp Arg Phe Ala Ser Leu Gln
520
Ser Leu Leu Leu Tyr Gln Leu Leu Gly Ile Phe His Arg Asp Glu
530
Gln Gln Arg Leu Leu Ser His Ser Phe His Ser Ala Leu Val Tyr
550
MET Phe Arg Ala Leu Asp Leu Pro Asn Arg Val Thr Gln Thr Ala
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Leu Val Lys Pro Arg Ala Asp MET His Gly Gly Glu Leu Glu Lys
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Ala Trp Lys Glu Trp Ile Glu Val Glu Thr Arg Arg Arg Val Ala
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Phe Ile Val Phe Leu Ile Asp Leu Glu His Ala Ala Ala Thr Glu
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Thr Pro Gln Leu Leu Ala Leu Ser Asp Leu Asp Leu Asp Leu Pro
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Ala Ser Glu Arg Ala Trp Lys Ala Glu Asn Ala Ala Glu Trp Leu
640
Asp Arg Ser Thr Ser Pro Leu Tyr Pro Glu Thr Ile Ser Phe Leu
650
Ala Ala Ile Arg Ala Leu Leu Ser Thr Thr Pro Pro Glu Pro Phe
670
Ser Ala Ala Gly Val Leu Leu Ala Glu Leu Gly Arg Leu Ser Ser
680
Phe Pro Leu Leu Ile Leu Ser Arg Thr Leu Ser Tyr Leu Glu Arg
700
Lys Thr Gln Glu Ala Leu Ala Gln Ile Asp Pro Phe Lys Ser Leu
710
Leu Gly Gly Leu Gly Val Leu Glu Gly Arg Glu Thr Glu Asn Arg
730
Ala Val Leu Glu Arg Ile Arg Arg Gly Arg Glu Val Leu Arg Arg
740
Leu Pro Gly Gly Ile Ala Arg Gly Gly Gly Glu Gly Trp Phe Asn
760
Glu Ile Ile Pro Ser Ala Lys Asp Phe Val Pro Ala Ser Asp Ser
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Pro Ala Ala Ser Ser Ser Ser Ala Gly Thr Ser Pro Arg Asp Ala
790
Phe Ser Ser Ser Ser Ala Ser Gly Thr Ala Ser Thr Phe Pro Ser
800
Pro Gln Asp Ala Pro Ala Ser Leu Glu Glu Leu Phe Ala Glu Phe
820
Asp Glu Gln Pro Tyr Arg Pro Phe His Gly Ala Gly Gly Ala Thr
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Pro Gly Glu Thr Tyr Glu Gln Ala Gln Glu Arg Leu Arg Lys His
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Trp Ala MET Pro ***
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<210> 3
<211> 27
<212> DNA
<213> 人工序列
<400> 3
aatggatcca tggcgcctgc tccccgt 27
<210> 4
<211> 29
<212> DNA
<213> 人工序列
<400> 4
atcgatatct cacggcatcg cccacacct 29
Claims (2)
1.一种锌指蛋白转录因子基因RkMSN4，其核苷酸序列如SEQ ID NO：1所示，该基因编码的氨基酸序列如SEQ ID NO：2所示。
2.权利要求1所述的锌指蛋白转录因子基因RkMSN4在低温下生产多不饱和脂肪酸中的应用。
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| CN109777815A (zh) * | 2019-03-28 | 2019-05-21 | 昆明理工大学 | HMG-CoA合成酶基因RKHMGCS及其应用 |
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