长囊水云与海带甘露醇代谢途径相关基因的研究
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摘要
甘露醇和海带淀粉是褐藻主要的碳积累产物，研究甘露醇代谢途径及其相关基因的调控变化，对高甘露醇含量的海带定向选育及遗传改良具有重要意义。
本研究根据褐藻长囊水云（E. siliculosus）的基因组数据，完成了2种甘露醇-1-磷酸酶（EsM1Pase1和EsM1Pase2）基因的结构及功能验证。通过克隆EsM1Pases基因的全长序列，进行了M1Pases的三维建模解析及其在不同藻类中的直系同源分析。经体外表达与组氨酸标签纯化，获得高纯度EsM1Pase2，发现其特异性的以甘露醇-1-磷酸（M1P）为底物，催化甘露醇的合成，而NaCl可以抑制该酶活性。通过测定不同M1P浓度下EsM1Pase2的反应速率，计算了该酶Km和Vm等动力学参数。通过qPCR检测，发现EsM1Pases基因表达具有明显的昼夜节律性，与甘露醇含量的变化规律一致。分析表明，长囊水云的M1Pases属于卤酸脱卤酶超家族（HAD super-family）的新成员。
结合长囊水云基因组及海带（S. japonica）转录组数据，获得海带甘露醇-2-脱氢酶（M2DH）的基因全长。qPCR检测结果表明，M2DH基因随海水盐度的降低，表达量增高；而在高浓度NaCl条件下，该基因几乎不转录，表明其对于低盐刺激具有较强的耐受性。在0.2-0.8mM H2O2处理及短时（0-2h）干露条件下，M2DH基因表达量升高，推测海带通过合成更多甘露醇来应答胁迫。2-6h的白光及蓝光照射组，M2DH基因表达量均高于黑暗组，而8-10h的黑暗处理略微上调该基因表达，表明不同光质诱导下，M2DH基因可能通过催化可逆反应的不同方向来调节甘露醇的含量。通过检测，甘露醇含量在不同品系、季节和部位的海带中均有差异。
本研究为解析褐藻甘露醇代谢途径与甘露醇积累的关系提供理论依据，也为含高甘露醇海带的选育及遗传改良提供参考。
Mannitol and laminarin are the main forms of carbon storage substances inbrown algae. Exploration of pathways and ralative genes involved with mannitolmetabolism is important for objective breeding and genetic improvement ofSaccharina with high-content mannitol.
Based on genomic data of Ectocarpus siliculosus, two genes encoding formannitol-1-phosphatase (named EsM1Pase1and EsM1Pase2) were isolated and theenzymatic function was verified subsequently. Through the amplification offull-length sequences, we analyzed the taxonomic distribution of EsM1Pase orthologsand spatial structure of EsM1Pase2. Recombinant EsM1Pase2was obtained via invitro expression in E. coli and purification by Ni2+affinity chromatography. It isshowed that EsM1Pase2was highly specific to hydrolyze mannitol-1-phosphate toproduce mannitol, and very sensitive to NaCl. Moreover, the kinetic parameters (Km,Vm etc.) were calculated with measurement of reaction velocity under various M1Pconcentrations. qPCR results showed that EsM1Pase2gene exhibited an analogousdiurnal rhythmicity with mannitol content. Our results demonstrated that M1Pase in E.siliculosus could be a new member of HAD super-family.
Combined E. siliculosus genome data with transcriptomic analysis of Saccharinajaponica, the full-length sequence of gene encoding for mannitol-2-dehydrogenase(M2DH) was obtained. qPCR detection for juvenile sporophytes showed that withsalinities decreasing, M2DH gene expression levels increased; while it was barelytranscribed under high NaCl concentrations, indicating M2DH exhibited strongtolerance to low salinities. Treatments with H2O2(0.2-0.8mM) and short-perioddessication (<2h) could up-regulate M2DH gene transcriptional levels, indicating thatthe kelp could produce more mannitol for abiotic stresss tolerance. Compared todarkness treatment, Sj-M2DH showed relatively higher expression levels under white and blue light for2-6h. When immersed in darkness for8-10h, M2DH transcriptsexhibited slight up-regulation. It was deduced that M2DH gene could regulatereversible enzymatic reaction to adjust mannitol content. In addition, the detectedmannitol content varied with different strains, seasons and positions of the kelp.
In conclusion, our study provided molecular data for deciphering the relationbetween mannitol pathway and its variations, and enriched genetic improvement ofthe kelp, which facilitated strains screening for large quantities of mannitol.
本研究根据褐藻长囊水云（E. siliculosus）的基因组数据，完成了2种甘露醇-1-磷酸酶（EsM1Pase1和EsM1Pase2）基因的结构及功能验证。通过克隆EsM1Pases基因的全长序列，进行了M1Pases的三维建模解析及其在不同藻类中的直系同源分析。经体外表达与组氨酸标签纯化，获得高纯度EsM1Pase2，发现其特异性的以甘露醇-1-磷酸（M1P）为底物，催化甘露醇的合成，而NaCl可以抑制该酶活性。通过测定不同M1P浓度下EsM1Pase2的反应速率，计算了该酶Km和Vm等动力学参数。通过qPCR检测，发现EsM1Pases基因表达具有明显的昼夜节律性，与甘露醇含量的变化规律一致。分析表明，长囊水云的M1Pases属于卤酸脱卤酶超家族（HAD super-family）的新成员。
结合长囊水云基因组及海带（S. japonica）转录组数据，获得海带甘露醇-2-脱氢酶（M2DH）的基因全长。qPCR检测结果表明，M2DH基因随海水盐度的降低，表达量增高；而在高浓度NaCl条件下，该基因几乎不转录，表明其对于低盐刺激具有较强的耐受性。在0.2-0.8mM H2O2处理及短时（0-2h）干露条件下，M2DH基因表达量升高，推测海带通过合成更多甘露醇来应答胁迫。2-6h的白光及蓝光照射组，M2DH基因表达量均高于黑暗组，而8-10h的黑暗处理略微上调该基因表达，表明不同光质诱导下，M2DH基因可能通过催化可逆反应的不同方向来调节甘露醇的含量。通过检测，甘露醇含量在不同品系、季节和部位的海带中均有差异。
本研究为解析褐藻甘露醇代谢途径与甘露醇积累的关系提供理论依据，也为含高甘露醇海带的选育及遗传改良提供参考。
Mannitol and laminarin are the main forms of carbon storage substances inbrown algae. Exploration of pathways and ralative genes involved with mannitolmetabolism is important for objective breeding and genetic improvement ofSaccharina with high-content mannitol.
Based on genomic data of Ectocarpus siliculosus, two genes encoding formannitol-1-phosphatase (named EsM1Pase1and EsM1Pase2) were isolated and theenzymatic function was verified subsequently. Through the amplification offull-length sequences, we analyzed the taxonomic distribution of EsM1Pase orthologsand spatial structure of EsM1Pase2. Recombinant EsM1Pase2was obtained via invitro expression in E. coli and purification by Ni2+affinity chromatography. It isshowed that EsM1Pase2was highly specific to hydrolyze mannitol-1-phosphate toproduce mannitol, and very sensitive to NaCl. Moreover, the kinetic parameters (Km,Vm etc.) were calculated with measurement of reaction velocity under various M1Pconcentrations. qPCR results showed that EsM1Pase2gene exhibited an analogousdiurnal rhythmicity with mannitol content. Our results demonstrated that M1Pase in E.siliculosus could be a new member of HAD super-family.
Combined E. siliculosus genome data with transcriptomic analysis of Saccharinajaponica, the full-length sequence of gene encoding for mannitol-2-dehydrogenase(M2DH) was obtained. qPCR detection for juvenile sporophytes showed that withsalinities decreasing, M2DH gene expression levels increased; while it was barelytranscribed under high NaCl concentrations, indicating M2DH exhibited strongtolerance to low salinities. Treatments with H2O2(0.2-0.8mM) and short-perioddessication (<2h) could up-regulate M2DH gene transcriptional levels, indicating thatthe kelp could produce more mannitol for abiotic stresss tolerance. Compared todarkness treatment, Sj-M2DH showed relatively higher expression levels under white and blue light for2-6h. When immersed in darkness for8-10h, M2DH transcriptsexhibited slight up-regulation. It was deduced that M2DH gene could regulatereversible enzymatic reaction to adjust mannitol content. In addition, the detectedmannitol content varied with different strains, seasons and positions of the kelp.
In conclusion, our study provided molecular data for deciphering the relationbetween mannitol pathway and its variations, and enriched genetic improvement ofthe kelp, which facilitated strains screening for large quantities of mannitol.
引文
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