基于香菇风味形成途径的内源性甲醛生成关键酶研究
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摘要
甲醛是香菇特征风味物质形成过程中的副产物之一,Y-谷氨酰转肽酶(GGT)和半胱氨酸亚砜裂解酶(C-S lyase)是风味形成途径中的关键酶。本研究旨在通过揭示关键酶对香菇内源性甲醛生成的作用来阐明风味形成途径对内源性甲醛形成的贡献。第一,分离纯化了香菇C-S lyase,并研究了该酶的蛋白质结构和部分酶学性质；第二,克隆得到香菇GGT基因全长cDNA序列,并研究了GGT基因mRNA表达水平的组织特异性、表达规律和不同储藏条件下的表达差异以及在相应条件下与甲醛生成之间的相关性。第三,通过添加外源物质和调整香菇培养基配方,直接或者间接调控GGT基因mRNA表达水平及其与C-S lyase酶活和特征风味物质的形成,量化了GGT和C-S lyase(?)寸香菇内源性甲醛生成的作用。主要研究结果如下：
1、C-S lyase的分离纯化及酶学性质的研究
(1)C-S lyase(?)勺快速分离纯化及其蛋白质结构
本研究主要通过两个关键步骤实现了对香菇中C-S lyase的快速分离和纯化,即先用PEG6000沉淀蛋白,然后经羟基磷灰石柱进-步分离,纯化倍数为234.82,得率为12.36%。纯化后的C-S lyase变性琼脂糖凝胶电泳(SDS-PAGE)、非变性琼脂糖凝胶电泳(Native-PAGE)和高效液相色谱(HPLC)研究结果表明,分离得到的C-S lyase具有较高纯度,同时该酶是由两个分子量约为44KDa的亚基组成的二聚体。通过氨基酸分析发现,C-S lyase由17种氨基酸组成,甲硫氨酸、丙氨酸和苯丙氨酸未被检出,酪氨酸含量最高(55%),其中酸性氨基酸与碱性氨基酸的比例为13.06/3.14。
(2)C-S lyase酶学性质的研究
酶学性质的研究结果表明,C-S lyase的等电点在pH4.75左右,最适反应pH和温度分别在7.5左右和35℃左右,并且具有较好的热稳定性,该酶对S-甲基-L-半胱氨酸亚砜的催化效率显著高于它对S-丙烯基-L-半胱氨酸亚砜和S-甲基-D-半胱氨酸亚砜的催化效率。此外,Cys和GSH以及Zn2+等金属离子对该酶的活性有抑制作用,而Lys则对酶活有促进作用。2、香菇GGT基因cDNA全长克隆及其mRNA表达水平
(1)香菇GGT基因cDNA全长克隆及序列分析
通过RACE技术克隆得到香菇GGT基因cDNA序列全长共2063bp (Gene bank登录号为JX123478),包括5'非翻译区序列36bp,开放读码框(ORF)序列1914bp,3'非翻译区序列113bp。ORF序列分析结果表明,该序列可编码由637个氨基酸残基组成的蛋白质,分子量约为70KDa,并且.含有GGT基因家族的保守核心结构域。BLAST结果表明,GGT氨基酸序列与裂褶菌、玉米丝黑穗病菌、皱木耳的同源性分别为94%、94%、92%。
(2) GGT基因在香菇和长过程、不同部位、不同储藏温度下的mRNA表达水平
生长过程研究结果表明,GGT基因mRNA表达水平随着香菇子实体的成熟先升高后降低,在第三个生长阶段达到最高表达量；不同部位的研究结果表明,菇柄中GGT mRNA的水平显著高于菇盖中的水平；不同品种研究结果表明,GGT基因在品种806中的mRNA表达水平高于在品种808中的表达水平；不同温度(4℃、20℃、30℃)储藏香菇的研究表明,GGT基因的mRNA表达量随着储藏温度的升高而降低；通过测定相应条件下甲醛的含量发现,GGT基因mRNA的表达与甲醛的生成规律基本一致。
3、风味形成途径对香菇内源性甲醛生成的影响
(1)直接添加从香菇中纯化得到的GGT和C-S lyase能显著提高香菇匀浆中甲醛的生成量；通过丝氨酸-硼酸复合物、GSH、Cys以及金属离子(Na+、K+、Mg2+、Zn2+、)抑制或者促进GGT和C-S lyase的酶活,匀浆中甲醛的生成量也显著的降低或者提高；改变香菇匀浆的酸碱度,甲醛的生成量随着随之改变,在pH8.0左右达到最大值,并且在弱碱性环境下(pH7-9)的甲醛生成量高于在弱酸性条件下(pH5-7)的生成量。
(2)通过改变香菇培养基中含硫物质(MgS04)的含量(0.1%、0.2%、0.4%)发现,香菇中甲醛的生成量随着含硫物质的增加而提高,同时发现香菇特征风味物质(1,2,4,5-四硫杂环己烷、香菇精)以及特征风味物质前体(S-丙烯基-L-半胱氨酸亚砜、S-甲基-L-半胱氨酸亚砜)、GGT mRNA水平和酶活以及C-S lyase的酶活的变化均与甲醛的生成规律基本一致。
综上所述,能够调控GGT基因表达水平、GGT和C-S lyase酶活、以及特征风味的上升或者下降的因素均能引起甲醛生成量的相应增加或者减少,表明香菇风味形成途径是内源性甲醛生成的主要途径。
Formaldehyde is one of by-products of odor-formed pathway where GGT and C-S lyase act as key enzymes. The aim of this study is to understand the contribution of odor-formed pathway to endogenous formaldehyde formation in shiitake mushroom (Lentinus edodes) by regulating activity or expression of GGT and C-S lyase. First, C-S lyase was purified from shiitake mushroom and its structures and properties were studied; second, the full-length cDNA sequence of GGT was cloned, and the relationship of GGT gene transcription and fonnaldehyde production in shiitake mushroom was discovered; third, effects of GGT and C-S lyase on the formaldehyde formation were studied by promoting or inhibiting activity of these two enzymes with their promoters and inhibitors addition into shiitake homogenate. Effects of MgSO4in medium on characteristic odors fonnation, odor precursors formation, formaldehyde formation, GGT gene expression, GGT and C-S lyase activity were studied. All results are shown as follows.
1Rapid purification and characterization of C-S lyase in shiitake mushroom
(1) Rapid purification and structures of C-S lyase
C-S lyase was purified from shiitake mushroom through two key steps, protein precipitation by PEG6000and C-S lyase elution by hydroxyapatite column, reaching to234.82purified fold and12.36%recovery. These results of SDS-PAGE, Native-PAGE and HPLC assays suggest the high purity of C-S lyase which consists of two identical subunits of44KDa. Amino acid composition of C-S lyase was analyzed, and17kinds of amino acids were detected. Of these amino acids, tyrosine accounts the highest with55%, and the ratio of the acidic amino acid and basic amino acid is13.06/3.14.
(2) Enzyme properties of C-S lyase
The result of isoelectric focusing electrophoresis (IEF) assay shows that the isoelectric point of C-S lyase is at pH4.75. The optimal temperature and pH value for enzymatic catalyzation are around35℃and7.5, respectively. C-S lyase is of good thermal stability. The catalytic efficiency of C-S lyase acting on S-methyl-L-cysteine sulfoxide is significantly higher than that acting on S-allyl-L-cysteine sulfoxide and S-methyl-D-cysteine sulfoxide. In addition, C-S lyase activity can be inhibited by Cys, GSH, and Zn2+, and be promoted by Lys.
2Cloning and analysis of GGT cDNA full-length sequence
(1) Cloning of GGT cDNA full-length sequence
The full-length cDNA sequence of GGT was obtained by RACE technology. The full-length cDNA sequence of GGT is2063bp (Gene bank accession NO. JX123478), including36bp of5'-untranslated region (UTR),113bp of3'-UTR and1914bp of open reading frame (ORF) that encodes a637-amino acid protein. The calculated molecular mass of this protein is70KDa, contained a GGT familial conserved domain. The BLAST analyasis of the deduced amino acid sequences of GGT with those of other species indicated that shiitake mushroom GGT sharing94%similarity with Schizophyllum commune,94%with Sporisorium reiliaimm,92%with Auricularia delicate.
(2) Expression pattern of GGT mRNA in shiitake mushroom
The developmental expression pattern of different growth stage study results show that in stalks and caps, GGT gene expression increases from the first stage to the third stage and then decreases significantly. Tissue-specific study results show that the GGT mRNA levels are higher expressed in stalks than in caps significantly. Strains study results show that GGT mRNA levels are higher in806than in808. Stored temperatures (4℃,20℃,30℃) study results show that the GGT mRNA level is the highest in shiitake mushroom stored at4℃and the lowest at30℃. Correspondingly, formaldehyde content was determined in growth process of shiitake mushroom. All these results indicate that the GGT gene expression positively affect formaldehyde formation.
3The contribution of odor-formed pathway to fonnaldehyde fonnation
(1) Effect of GGT and C-S lyase on formaldehyde fonnation
To homogenate of shiitake mushroom, adding purified GGT and C-S lyase, Na+and K+can promote the fonnaldehyde fonnation, and adding Ser-borate, GSH, Cys and Zn2+can inhibit the fonnaldehyde fonnation. In addition, as regulating the homogenate pH value from4to10, the formaldehyde formation increase gradually until reaches the highest at pH8and then decreases from pH9to10.
(2) Effect of MgSO4in medium on fonnaldehyde and odor fonnation
Of shiitake mushrooms cultured in medium with MgSO4level of0.1%,0.2%,0.4%, respectively, formaldehyde, odor compounds (1,2,4,5-Tetrathiane and Lenthionine), odor precursors (S-allyl-L-cysteine sulfoxide and S-methyl-L-cysteine sulfoxide), GGT mRNA level, GGT activity, and C-S lyase activity were measured. Results show that, as MgSO4content increases the fonnation of fonnaldehyde, odor compounds and odor precursors rises following the enhance of GGT gene expression and enzyme activity of GGT and C-S lyase.
As a conclusion, the odor-formed pathway is the main pathway of fonnaldehyde fonnation in shiitake mushroom.
1、C-S lyase的分离纯化及酶学性质的研究
(1)C-S lyase(?)勺快速分离纯化及其蛋白质结构
本研究主要通过两个关键步骤实现了对香菇中C-S lyase的快速分离和纯化,即先用PEG6000沉淀蛋白,然后经羟基磷灰石柱进-步分离,纯化倍数为234.82,得率为12.36%。纯化后的C-S lyase变性琼脂糖凝胶电泳(SDS-PAGE)、非变性琼脂糖凝胶电泳(Native-PAGE)和高效液相色谱(HPLC)研究结果表明,分离得到的C-S lyase具有较高纯度,同时该酶是由两个分子量约为44KDa的亚基组成的二聚体。通过氨基酸分析发现,C-S lyase由17种氨基酸组成,甲硫氨酸、丙氨酸和苯丙氨酸未被检出,酪氨酸含量最高(55%),其中酸性氨基酸与碱性氨基酸的比例为13.06/3.14。
(2)C-S lyase酶学性质的研究
酶学性质的研究结果表明,C-S lyase的等电点在pH4.75左右,最适反应pH和温度分别在7.5左右和35℃左右,并且具有较好的热稳定性,该酶对S-甲基-L-半胱氨酸亚砜的催化效率显著高于它对S-丙烯基-L-半胱氨酸亚砜和S-甲基-D-半胱氨酸亚砜的催化效率。此外,Cys和GSH以及Zn2+等金属离子对该酶的活性有抑制作用,而Lys则对酶活有促进作用。2、香菇GGT基因cDNA全长克隆及其mRNA表达水平
(1)香菇GGT基因cDNA全长克隆及序列分析
通过RACE技术克隆得到香菇GGT基因cDNA序列全长共2063bp (Gene bank登录号为JX123478),包括5'非翻译区序列36bp,开放读码框(ORF)序列1914bp,3'非翻译区序列113bp。ORF序列分析结果表明,该序列可编码由637个氨基酸残基组成的蛋白质,分子量约为70KDa,并且.含有GGT基因家族的保守核心结构域。BLAST结果表明,GGT氨基酸序列与裂褶菌、玉米丝黑穗病菌、皱木耳的同源性分别为94%、94%、92%。
(2) GGT基因在香菇和长过程、不同部位、不同储藏温度下的mRNA表达水平
生长过程研究结果表明,GGT基因mRNA表达水平随着香菇子实体的成熟先升高后降低,在第三个生长阶段达到最高表达量；不同部位的研究结果表明,菇柄中GGT mRNA的水平显著高于菇盖中的水平；不同品种研究结果表明,GGT基因在品种806中的mRNA表达水平高于在品种808中的表达水平；不同温度(4℃、20℃、30℃)储藏香菇的研究表明,GGT基因的mRNA表达量随着储藏温度的升高而降低；通过测定相应条件下甲醛的含量发现,GGT基因mRNA的表达与甲醛的生成规律基本一致。
3、风味形成途径对香菇内源性甲醛生成的影响
(1)直接添加从香菇中纯化得到的GGT和C-S lyase能显著提高香菇匀浆中甲醛的生成量；通过丝氨酸-硼酸复合物、GSH、Cys以及金属离子(Na+、K+、Mg2+、Zn2+、)抑制或者促进GGT和C-S lyase的酶活,匀浆中甲醛的生成量也显著的降低或者提高；改变香菇匀浆的酸碱度,甲醛的生成量随着随之改变,在pH8.0左右达到最大值,并且在弱碱性环境下(pH7-9)的甲醛生成量高于在弱酸性条件下(pH5-7)的生成量。
(2)通过改变香菇培养基中含硫物质(MgS04)的含量(0.1%、0.2%、0.4%)发现,香菇中甲醛的生成量随着含硫物质的增加而提高,同时发现香菇特征风味物质(1,2,4,5-四硫杂环己烷、香菇精)以及特征风味物质前体(S-丙烯基-L-半胱氨酸亚砜、S-甲基-L-半胱氨酸亚砜)、GGT mRNA水平和酶活以及C-S lyase的酶活的变化均与甲醛的生成规律基本一致。
综上所述,能够调控GGT基因表达水平、GGT和C-S lyase酶活、以及特征风味的上升或者下降的因素均能引起甲醛生成量的相应增加或者减少,表明香菇风味形成途径是内源性甲醛生成的主要途径。
Formaldehyde is one of by-products of odor-formed pathway where GGT and C-S lyase act as key enzymes. The aim of this study is to understand the contribution of odor-formed pathway to endogenous formaldehyde formation in shiitake mushroom (Lentinus edodes) by regulating activity or expression of GGT and C-S lyase. First, C-S lyase was purified from shiitake mushroom and its structures and properties were studied; second, the full-length cDNA sequence of GGT was cloned, and the relationship of GGT gene transcription and fonnaldehyde production in shiitake mushroom was discovered; third, effects of GGT and C-S lyase on the formaldehyde formation were studied by promoting or inhibiting activity of these two enzymes with their promoters and inhibitors addition into shiitake homogenate. Effects of MgSO4in medium on characteristic odors fonnation, odor precursors formation, formaldehyde formation, GGT gene expression, GGT and C-S lyase activity were studied. All results are shown as follows.
1Rapid purification and characterization of C-S lyase in shiitake mushroom
(1) Rapid purification and structures of C-S lyase
C-S lyase was purified from shiitake mushroom through two key steps, protein precipitation by PEG6000and C-S lyase elution by hydroxyapatite column, reaching to234.82purified fold and12.36%recovery. These results of SDS-PAGE, Native-PAGE and HPLC assays suggest the high purity of C-S lyase which consists of two identical subunits of44KDa. Amino acid composition of C-S lyase was analyzed, and17kinds of amino acids were detected. Of these amino acids, tyrosine accounts the highest with55%, and the ratio of the acidic amino acid and basic amino acid is13.06/3.14.
(2) Enzyme properties of C-S lyase
The result of isoelectric focusing electrophoresis (IEF) assay shows that the isoelectric point of C-S lyase is at pH4.75. The optimal temperature and pH value for enzymatic catalyzation are around35℃and7.5, respectively. C-S lyase is of good thermal stability. The catalytic efficiency of C-S lyase acting on S-methyl-L-cysteine sulfoxide is significantly higher than that acting on S-allyl-L-cysteine sulfoxide and S-methyl-D-cysteine sulfoxide. In addition, C-S lyase activity can be inhibited by Cys, GSH, and Zn2+, and be promoted by Lys.
2Cloning and analysis of GGT cDNA full-length sequence
(1) Cloning of GGT cDNA full-length sequence
The full-length cDNA sequence of GGT was obtained by RACE technology. The full-length cDNA sequence of GGT is2063bp (Gene bank accession NO. JX123478), including36bp of5'-untranslated region (UTR),113bp of3'-UTR and1914bp of open reading frame (ORF) that encodes a637-amino acid protein. The calculated molecular mass of this protein is70KDa, contained a GGT familial conserved domain. The BLAST analyasis of the deduced amino acid sequences of GGT with those of other species indicated that shiitake mushroom GGT sharing94%similarity with Schizophyllum commune,94%with Sporisorium reiliaimm,92%with Auricularia delicate.
(2) Expression pattern of GGT mRNA in shiitake mushroom
The developmental expression pattern of different growth stage study results show that in stalks and caps, GGT gene expression increases from the first stage to the third stage and then decreases significantly. Tissue-specific study results show that the GGT mRNA levels are higher expressed in stalks than in caps significantly. Strains study results show that GGT mRNA levels are higher in806than in808. Stored temperatures (4℃,20℃,30℃) study results show that the GGT mRNA level is the highest in shiitake mushroom stored at4℃and the lowest at30℃. Correspondingly, formaldehyde content was determined in growth process of shiitake mushroom. All these results indicate that the GGT gene expression positively affect formaldehyde formation.
3The contribution of odor-formed pathway to fonnaldehyde fonnation
(1) Effect of GGT and C-S lyase on formaldehyde fonnation
To homogenate of shiitake mushroom, adding purified GGT and C-S lyase, Na+and K+can promote the fonnaldehyde fonnation, and adding Ser-borate, GSH, Cys and Zn2+can inhibit the fonnaldehyde fonnation. In addition, as regulating the homogenate pH value from4to10, the formaldehyde formation increase gradually until reaches the highest at pH8and then decreases from pH9to10.
(2) Effect of MgSO4in medium on fonnaldehyde and odor fonnation
Of shiitake mushrooms cultured in medium with MgSO4level of0.1%,0.2%,0.4%, respectively, formaldehyde, odor compounds (1,2,4,5-Tetrathiane and Lenthionine), odor precursors (S-allyl-L-cysteine sulfoxide and S-methyl-L-cysteine sulfoxide), GGT mRNA level, GGT activity, and C-S lyase activity were measured. Results show that, as MgSO4content increases the fonnation of fonnaldehyde, odor compounds and odor precursors rises following the enhance of GGT gene expression and enzyme activity of GGT and C-S lyase.
As a conclusion, the odor-formed pathway is the main pathway of fonnaldehyde fonnation in shiitake mushroom.
引文
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