本文主要研究内容
作者邵璇（2019）在《海洋玫瑰杆菌类群细菌代谢二甲基巯基丙酸内盐脱甲基途径关键酶的催化机制及该途径的动力学调控机制》一文中研究指出：二甲基疏基丙酸内盐(Dimethylsulfoniopropionate,DMSP)是海洋碳硫循环的重要载体物质。每年由海洋浮游植物、大型藻类和细菌产生的DMSP的量可达上亿吨,可占部分海洋表面固碳总量的10%。DMSP具有多种不同的生理功能。做为作为一种重要的硫源和碳源,DMSP可以被不同种类的细菌利用,其中海洋玫瑰杆菌类群和SAR11类群是最主要的参与者。海洋细菌通过两个途径代谢DMSP,即裂解途径和脱甲基途径。其中,大约10%的DMSP经由裂解途径代谢,绝大多数DMSP通过脱甲基途径代谢。通过裂解途径,DMSP被多种裂解酶裂解产生二甲基硫和丙烯酸,丙烯酸被PrpE和AcuI进一步代谢,被细菌利用。在脱甲基途径中,DMSP在四个酶(DmdA、DmdB、DmdC和DmdD/AcuH)的作用下被分解代谢并生成甲硫醇,被细菌利用。脱甲基途径承担着绝大多数DMSP的代谢,因此研究该途径中关键酶的结构、催化机制以及在该途径中DMSP代谢的动力学调控机制,有助于更好的研究DMSP的分解代谢过程,并对研究海洋微生物活动对全球碳硫循环的推动作用具有重要意义。DMSP脱甲基途径中两个关键酶DmdA和DmdD/AcuH的结构和催化机制已被报道,但另外两个关键酶DmdB和DmdC的结构和催化机制还没有被报道。在本论文中,我们以海洋玫瑰杆菌类群细菌为研究对象,研究了DMSP脱甲基途径中DmdB和DmdC的结构和催化机制以及海洋玫瑰杆菌类群细菌中DMSP脱甲基代谢途径的动力学调控机制。论文取得了如下结果:一、海洋玫瑰杆菌类群细菌中MMPA-CoA连接酶DmdB的结构及其催化的分子机制海洋细菌DMSP脱甲基途径中的关键酶DmdB是一个3-甲基巯基丙酸(3-methylmercaptopropionate,MMPA)-辅酶A(Coenzyme,CoA)连接酶,催化DMSP脱甲基途径中脱甲基酶DmdA的产物MMPA与CoA的连接,并产生MMPA-CoA。我们以海洋玫瑰杆菌类群细菌Ruegeria lacuscaerulensis ITI-11157来源的DmdB为研究对象,研究了DmdB的结构和催化分子机制。我们首先利用RT-qPCR技术和酶活检测实验手段验证了dmd 基因的功能。实验结果表明dmdB基因能够被DMSP诱导转录上调,且重组表达纯化的DmdB具有显著的MMPA-CoA连接酶活性。然后我们分析了DmdB的酶学性质。凝胶过滤层析结果表明DmdB在溶液中以二聚体形式发挥功能。然后我们分别解析了DmdB结合ATP分子的竞争性抑制剂ADP以及DmdB突变体Lys523Ala结合AMP和MMPA的复合物晶体结构。从结构中可以看出,DmdB是一个二聚体,它每个单体包括一个大的N末端结构域和一个小的C末端结构域,且催化活性中心位于这两个结构域之间。DmdB在催化过程中会发生两次构象改变,当不结合任何配体时,DmdB处于开放构象,ATP分子的结合促使DmdB的C末端结构域发生～64°的旋转,使DmdB发生第一次构象改变,形成腺苷形成构象,并在此构象下DmdB催化第一步反应。利用序列及结构分析、定点突变实验验证和圆二色光谱分析,我们发现Lys523是重要的催化氨基酸残基,它通过与MMPA和ATP形成作用力来参与催化反应。然后,我们系统地研究了 DmdB催化中心多个保守氨基酸残基的功能,发现了与底物结合以及催化相关的氨基酸残基。通过将DmdB结构与已报道的其他辅酶A连接酶结构叠加,我们分析了DmdB的硫酯形成构象,以及在该构象下的关键氨基酸残基的功能。实验结果表明DmdB的C末端结构域会再次经过旋转使DmdB发生第二次构象变化,变为硫酯形成构象,并在此构象下DmdB催化第二步反应。最后,综合实验结果,我们提出了DmdB催化MMPA和CoA连接生成MMPA-CoA的分子机制。多序列比对分析表明,DmdB的催化机制可能在海洋细菌中具有普遍适用性。本研究对更好的认识海洋细菌通过脱甲基途径代谢DMSP的过程具有重要意义。二、海洋玫瑰杆菌类群细菌中MMPA-CoA脱氢酶DmdC的结构及其催化的分子机制在DMSP脱甲基途径上,MMPA-CoA脱氢酶DmdC利用FAD做为辅因子催化上游DmdB的产物MMPA-CoA生成3-甲基巯基丙烯酰辅酶A(methylthioacrylyl-CoA,MTA-CoA)。我们以海洋玫瑰杆菌类群细菌Roseovarius nubinhibens ISM来源的DmdC为研究对象,研究了DmdC的结构和催化机制。首先,通过RT-qPCR技术我们验证了菌株ISM中dmdC基因的功能。实验结果表明DMSP显著诱导dmdC基因转录上调,且重组表达纯化的DmdC具有显著的MMPA-CoA脱氢酶活性。然后我们分析了DmdC的酶学性质。在此基础上进一步解析了DmdC的晶体结构。结构和生化分析表明,DmdC是一个二聚体,它的两个单体通过一个大的接触表面组装在一起。通过分子对接研究,我们预测了DmdC的结构中FAD的结合位点以及催化中心一些关键氨基酸残基的功能;然后结合定点突变实验和圆二色光谱分析结果,我们确定了DmdC中与FAD结合及与催化相关的一些氨基酸残基。通过将DmdC结构与同家族已报道的结合了底物分子的其他酰基辅酶A脱氢酶的结构的叠加,我们分析了DmdC的结构中底物MMPA-CoA的结合位点,并结合定点突变实验结果确定了与底物MMPA-CoA结合相关的氨基酸残基以及参与催化反应的催化氨基酸残基。最后,综合实验结果,我们提出了DmdC催化MMPA-CoA脱氢生成MTA-CoA的分子机制。通过序列比对发现该机制可能在不同细菌来源的DmdC中普遍适用。本研究对更全面的认识海洋细菌的DMSP脱甲基代谢途径具有重要意义。三、海洋玫瑰杆菌类群细菌DMSP脱甲基代谢途径的动力学调控机制在海洋细菌中,DMSP的分解代谢是一个复杂的过程。海洋细菌通过裂解途径和脱甲基途径中的多个酶的协同作用完成DMSP的分解代谢并同时保证DMSP的生理功能。海水中的DMSP浓度很低,为纳摩尔级别,而经过富集,在海洋细菌胞内其浓度可达到毫摩尔量级。已经有研究人员揭示在裂解途径上的DMSP分解代谢的动力学调控机制。本论文研究了 DMSP分解代谢的脱甲基途径的动力学调控机制。通过RT-qPCR和胞外酶活,我们首先证实了海洋玫瑰杆菌类群细菌中DMSP脱甲基途径上的四个酶基因的功能。然后通过测定酶的Km值分析了这四个酶对底物的亲和力。结果表明,不同菌株来源的DMSP脱甲基酶DmdA对底物DMSP的Km值为十个毫摩尔量级,和DMSP裂解酶一样。这表明DmdA对底物DMSP的亲和力较低,这可以保证DMSP在海洋细菌胞内维持较高的浓度以发挥其生理功能。DmdB和DmdC两个酶对底物的Km值与DMSP裂解途径中的丙酸辅酶A连接酶PrpE的Km值处于相同的水平,都在毫摩尔水平且低于DmdA的Km值,这表明DmdB和DmdC对底物的亲和力高于DmdA。底物亲和力的提高,保证了 DMSP的继续代谢,有利于物质转化和能量流动。DMSP脱甲基途径的最后一个酶为MTA-CoA水合酶DmdD,其对底物的Km值为微摩尔级别,且催化效率非常高。DmdD如此高的底物亲和力和催化效率,保证了底物的快速代谢,避免其在细胞内的积累,这与已报道的AcuI相同,也从侧面上暗示了底物MTA-CoA可能具有细胞毒害作用。基于我们的分析,我们提出了玫瑰杆菌类群细菌中DMSP脱甲基途径的动力学调控机制,即参与DMSP脱甲基代谢的四个酶DmdA、DmdB、DmdC和DmdD通过对底物亲和力的调控作用,保证了DMSP的胞内积累,以维持其生理功能;同时还保证了DMSP及其代谢产物尤其是有毒的MTA-CoA的快速代谢,最终完成DMSP的脱甲基代谢过程。本论文对海洋玫瑰杆菌类群细菌DMSP脱甲基代谢途径中关键酶DmdB和DmdC的晶体结构、催化机制和DMSP脱甲基代谢途径的动力学调控机制进行了较为深入的研究,研究结果有助于我们更好的了解海洋细菌对DMSP的分解代谢过程,以及更好的认识海洋碳、硫循环。四、论文的其他研究结果:深海细菌Myroides profundi D25产弹性蛋白酶myroilysin的小试和中试发酵条件优化在系统研究海洋细菌代谢重要有机硫DMSP机制的基础上,本论文还开展了一株深海沉积物来源细菌产弹性蛋白酶的发酵工艺研究。前期研究表明,弹性蛋白酶myroilysin是深海沉积物来源细菌Myroides profundi D25分泌的主要蛋白酶,它同时具有显著的膨胀胶原蛋白的能力,这表明myroilysin在生物技术应用方面具有很大潜力。由于myroilysin不能自体成熟,因此提高野生M.profundi D25细菌中myroilysin的产量尤其重要。在本论文中,我们首先利用单因子实验优化了菌株Mrofundi D25的培养条件。随后,利用优化后的培养条件,我们进行了多次菌株M.profundi D25的小试和中试发酵,并成功建立了M.profundi D25产myroilysin的小试和中试发酵工艺。发酵工艺的建立,为蛋白酶myroilysin的工业化生产和其生物技术方面的开发应用奠定了基础。
Abstract
er jia ji shu ji bing suan nei yan (Dimethylsulfoniopropionate,DMSP)shi hai xiang tan liu xun huan de chong yao zai ti wu zhi 。mei nian you hai xiang fu you zhi wu 、da xing zao lei he xi jun chan sheng de DMSPde liang ke da shang yi dun ,ke zhan bu fen hai xiang biao mian gu tan zong liang de 10%。DMSPju you duo chong bu tong de sheng li gong neng 。zuo wei zuo wei yi chong chong yao de liu yuan he tan yuan ,DMSPke yi bei bu tong chong lei de xi jun li yong ,ji zhong hai xiang mei gui gan jun lei qun he SAR11lei qun shi zui zhu yao de can yu zhe 。hai xiang xi jun tong guo liang ge tu jing dai xie DMSP,ji lie jie tu jing he tuo jia ji tu jing 。ji zhong ,da yao 10%de DMSPjing you lie jie tu jing dai xie ,jue da duo shu DMSPtong guo tuo jia ji tu jing dai xie 。tong guo lie jie tu jing ,DMSPbei duo chong lie jie mei lie jie chan sheng er jia ji liu he bing xi suan ,bing xi suan bei PrpEhe AcuIjin yi bu dai xie ,bei xi jun li yong 。zai tuo jia ji tu jing zhong ,DMSPzai si ge mei (DmdA、DmdB、DmdChe DmdD/AcuH)de zuo yong xia bei fen jie dai xie bing sheng cheng jia liu chun ,bei xi jun li yong 。tuo jia ji tu jing cheng dan zhao jue da duo shu DMSPde dai xie ,yin ci yan jiu gai tu jing zhong guan jian mei de jie gou 、cui hua ji zhi yi ji zai gai tu jing zhong DMSPdai xie de dong li xue diao kong ji zhi ,you zhu yu geng hao de yan jiu DMSPde fen jie dai xie guo cheng ,bing dui yan jiu hai xiang wei sheng wu huo dong dui quan qiu tan liu xun huan de tui dong zuo yong ju you chong yao yi yi 。DMSPtuo jia ji tu jing zhong liang ge guan jian mei DmdAhe DmdD/AcuHde jie gou he cui hua ji zhi yi bei bao dao ,dan ling wai liang ge guan jian mei DmdBhe DmdCde jie gou he cui hua ji zhi hai mei you bei bao dao 。zai ben lun wen zhong ,wo men yi hai xiang mei gui gan jun lei qun xi jun wei yan jiu dui xiang ,yan jiu le DMSPtuo jia ji tu jing zhong DmdBhe DmdCde jie gou he cui hua ji zhi yi ji hai xiang mei gui gan jun lei qun xi jun zhong DMSPtuo jia ji dai xie tu jing de dong li xue diao kong ji zhi 。lun wen qu de le ru xia jie guo :yi 、hai xiang mei gui gan jun lei qun xi jun zhong MMPA-CoAlian jie mei DmdBde jie gou ji ji cui hua de fen zi ji zhi hai xiang xi jun DMSPtuo jia ji tu jing zhong de guan jian mei DmdBshi yi ge 3-jia ji qiu ji bing suan (3-methylmercaptopropionate,MMPA)-fu mei A(Coenzyme,CoA)lian jie mei ,cui hua DMSPtuo jia ji tu jing zhong tuo jia ji mei DmdAde chan wu MMPAyu CoAde lian jie ,bing chan sheng MMPA-CoA。wo men yi hai xiang mei gui gan jun lei qun xi jun Ruegeria lacuscaerulensis ITI-11157lai yuan de DmdBwei yan jiu dui xiang ,yan jiu le DmdBde jie gou he cui hua fen zi ji zhi 。wo men shou xian li yong RT-qPCRji shu he mei huo jian ce shi yan shou duan yan zheng le dmd ji yin de gong neng 。shi yan jie guo biao ming dmdBji yin neng gou bei DMSPyou dao zhuai lu shang diao ,ju chong zu biao da chun hua de DmdBju you xian zhe de MMPA-CoAlian jie mei huo xing 。ran hou wo men fen xi le DmdBde mei xue xing zhi 。ning jiao guo lv ceng xi jie guo biao ming DmdBzai rong ye zhong yi er ju ti xing shi fa hui gong neng 。ran hou wo men fen bie jie xi le DmdBjie ge ATPfen zi de jing zheng xing yi zhi ji ADPyi ji DmdBtu bian ti Lys523Alajie ge AMPhe MMPAde fu ge wu jing ti jie gou 。cong jie gou zhong ke yi kan chu ,DmdBshi yi ge er ju ti ,ta mei ge chan ti bao gua yi ge da de Nmo duan jie gou yu he yi ge xiao de Cmo duan jie gou yu ,ju cui hua huo xing zhong xin wei yu zhe liang ge jie gou yu zhi jian 。DmdBzai cui hua guo cheng zhong hui fa sheng liang ci gou xiang gai bian ,dang bu jie ge ren he pei ti shi ,DmdBchu yu kai fang gou xiang ,ATPfen zi de jie ge cu shi DmdBde Cmo duan jie gou yu fa sheng ～64°de xuan zhuai ,shi DmdBfa sheng di yi ci gou xiang gai bian ,xing cheng xian gan xing cheng gou xiang ,bing zai ci gou xiang xia DmdBcui hua di yi bu fan ying 。li yong xu lie ji jie gou fen xi 、ding dian tu bian shi yan yan zheng he yuan er se guang pu fen xi ,wo men fa xian Lys523shi chong yao de cui hua an ji suan can ji ,ta tong guo yu MMPAhe ATPxing cheng zuo yong li lai can yu cui hua fan ying 。ran hou ,wo men ji tong de yan jiu le DmdBcui hua zhong xin duo ge bao shou an ji suan can ji de gong neng ,fa xian le yu de wu jie ge yi ji cui hua xiang guan de an ji suan can ji 。tong guo jiang DmdBjie gou yu yi bao dao de ji ta fu mei Alian jie mei jie gou die jia ,wo men fen xi le DmdBde liu zhi xing cheng gou xiang ,yi ji zai gai gou xiang xia de guan jian an ji suan can ji de gong neng 。shi yan jie guo biao ming DmdBde Cmo duan jie gou yu hui zai ci jing guo xuan zhuai shi DmdBfa sheng di er ci gou xiang bian hua ,bian wei liu zhi xing cheng gou xiang ,bing zai ci gou xiang xia DmdBcui hua di er bu fan ying 。zui hou ,zeng ge shi yan jie guo ,wo men di chu le DmdBcui hua MMPAhe CoAlian jie sheng cheng MMPA-CoAde fen zi ji zhi 。duo xu lie bi dui fen xi biao ming ,DmdBde cui hua ji zhi ke neng zai hai xiang xi jun zhong ju you pu bian kuo yong xing 。ben yan jiu dui geng hao de ren shi hai xiang xi jun tong guo tuo jia ji tu jing dai xie DMSPde guo cheng ju you chong yao yi yi 。er 、hai xiang mei gui gan jun lei qun xi jun zhong MMPA-CoAtuo qing mei DmdCde jie gou ji ji cui hua de fen zi ji zhi zai DMSPtuo jia ji tu jing shang ,MMPA-CoAtuo qing mei DmdCli yong FADzuo wei fu yin zi cui hua shang you DmdBde chan wu MMPA-CoAsheng cheng 3-jia ji qiu ji bing xi xian fu mei A(methylthioacrylyl-CoA,MTA-CoA)。wo men yi hai xiang mei gui gan jun lei qun xi jun Roseovarius nubinhibens ISMlai yuan de DmdCwei yan jiu dui xiang ,yan jiu le DmdCde jie gou he cui hua ji zhi 。shou xian ,tong guo RT-qPCRji shu wo men yan zheng le jun zhu ISMzhong dmdCji yin de gong neng 。shi yan jie guo biao ming DMSPxian zhe you dao dmdCji yin zhuai lu shang diao ,ju chong zu biao da chun hua de DmdCju you xian zhe de MMPA-CoAtuo qing mei huo xing 。ran hou wo men fen xi le DmdCde mei xue xing zhi 。zai ci ji chu shang jin yi bu jie xi le DmdCde jing ti jie gou 。jie gou he sheng hua fen xi biao ming ,DmdCshi yi ge er ju ti ,ta de liang ge chan ti tong guo yi ge da de jie chu biao mian zu zhuang zai yi qi 。tong guo fen zi dui jie yan jiu ,wo men yu ce le DmdCde jie gou zhong FADde jie ge wei dian yi ji cui hua zhong xin yi xie guan jian an ji suan can ji de gong neng ;ran hou jie ge ding dian tu bian shi yan he yuan er se guang pu fen xi jie guo ,wo men que ding le DmdCzhong yu FADjie ge ji yu cui hua xiang guan de yi xie an ji suan can ji 。tong guo jiang DmdCjie gou yu tong jia zu yi bao dao de jie ge le de wu fen zi de ji ta xian ji fu mei Atuo qing mei de jie gou de die jia ,wo men fen xi le DmdCde jie gou zhong de wu MMPA-CoAde jie ge wei dian ,bing jie ge ding dian tu bian shi yan jie guo que ding le yu de wu MMPA-CoAjie ge xiang guan de an ji suan can ji yi ji can yu cui hua fan ying de cui hua an ji suan can ji 。zui hou ,zeng ge shi yan jie guo ,wo men di chu le DmdCcui hua MMPA-CoAtuo qing sheng cheng MTA-CoAde fen zi ji zhi 。tong guo xu lie bi dui fa xian gai ji zhi ke neng zai bu tong xi jun lai yuan de DmdCzhong pu bian kuo yong 。ben yan jiu dui geng quan mian de ren shi hai xiang xi jun de DMSPtuo jia ji dai xie tu jing ju you chong yao yi yi 。san 、hai xiang mei gui gan jun lei qun xi jun DMSPtuo jia ji dai xie tu jing de dong li xue diao kong ji zhi zai hai xiang xi jun zhong ,DMSPde fen jie dai xie shi yi ge fu za de guo cheng 。hai xiang xi jun tong guo lie jie tu jing he tuo jia ji tu jing zhong de duo ge mei de xie tong zuo yong wan cheng DMSPde fen jie dai xie bing tong shi bao zheng DMSPde sheng li gong neng 。hai shui zhong de DMSPnong du hen di ,wei na ma er ji bie ,er jing guo fu ji ,zai hai xiang xi jun bao nei ji nong du ke da dao hao ma er liang ji 。yi jing you yan jiu ren yuan jie shi zai lie jie tu jing shang de DMSPfen jie dai xie de dong li xue diao kong ji zhi 。ben lun wen yan jiu le DMSPfen jie dai xie de tuo jia ji tu jing de dong li xue diao kong ji zhi 。tong guo RT-qPCRhe bao wai mei huo ,wo men shou xian zheng shi le hai xiang mei gui gan jun lei qun xi jun zhong DMSPtuo jia ji tu jing shang de si ge mei ji yin de gong neng 。ran hou tong guo ce ding mei de Kmzhi fen xi le zhe si ge mei dui de wu de qin he li 。jie guo biao ming ,bu tong jun zhu lai yuan de DMSPtuo jia ji mei DmdAdui de wu DMSPde Kmzhi wei shi ge hao ma er liang ji ,he DMSPlie jie mei yi yang 。zhe biao ming DmdAdui de wu DMSPde qin he li jiao di ,zhe ke yi bao zheng DMSPzai hai xiang xi jun bao nei wei chi jiao gao de nong du yi fa hui ji sheng li gong neng 。DmdBhe DmdCliang ge mei dui de wu de Kmzhi yu DMSPlie jie tu jing zhong de bing suan fu mei Alian jie mei PrpEde Kmzhi chu yu xiang tong de shui ping ,dou zai hao ma er shui ping ju di yu DmdAde Kmzhi ,zhe biao ming DmdBhe DmdCdui de wu de qin he li gao yu DmdA。de wu qin he li de di gao ,bao zheng le DMSPde ji xu dai xie ,you li yu wu zhi zhuai hua he neng liang liu dong 。DMSPtuo jia ji tu jing de zui hou yi ge mei wei MTA-CoAshui ge mei DmdD,ji dui de wu de Kmzhi wei wei ma er ji bie ,ju cui hua xiao lv fei chang gao 。DmdDru ci gao de de wu qin he li he cui hua xiao lv ,bao zheng le de wu de kuai su dai xie ,bi mian ji zai xi bao nei de ji lei ,zhe yu yi bao dao de AcuIxiang tong ,ye cong ce mian shang an shi le de wu MTA-CoAke neng ju you xi bao du hai zuo yong 。ji yu wo men de fen xi ,wo men di chu le mei gui gan jun lei qun xi jun zhong DMSPtuo jia ji tu jing de dong li xue diao kong ji zhi ,ji can yu DMSPtuo jia ji dai xie de si ge mei DmdA、DmdB、DmdChe DmdDtong guo dui de wu qin he li de diao kong zuo yong ,bao zheng le DMSPde bao nei ji lei ,yi wei chi ji sheng li gong neng ;tong shi hai bao zheng le DMSPji ji dai xie chan wu you ji shi you du de MTA-CoAde kuai su dai xie ,zui zhong wan cheng DMSPde tuo jia ji dai xie guo cheng 。ben lun wen dui hai xiang mei gui gan jun lei qun xi jun DMSPtuo jia ji dai xie tu jing zhong guan jian mei DmdBhe DmdCde jing ti jie gou 、cui hua ji zhi he DMSPtuo jia ji dai xie tu jing de dong li xue diao kong ji zhi jin hang le jiao wei shen ru de yan jiu ,yan jiu jie guo you zhu yu wo men geng hao de le jie hai xiang xi jun dui DMSPde fen jie dai xie guo cheng ,yi ji geng hao de ren shi hai xiang tan 、liu xun huan 。si 、lun wen de ji ta yan jiu jie guo :shen hai xi jun Myroides profundi D25chan dan xing dan bai mei myroilysinde xiao shi he zhong shi fa jiao tiao jian you hua zai ji tong yan jiu hai xiang xi jun dai xie chong yao you ji liu DMSPji zhi de ji chu shang ,ben lun wen hai kai zhan le yi zhu shen hai chen ji wu lai yuan xi jun chan dan xing dan bai mei de fa jiao gong yi yan jiu 。qian ji yan jiu biao ming ,dan xing dan bai mei myroilysinshi shen hai chen ji wu lai yuan xi jun Myroides profundi D25fen bi de zhu yao dan bai mei ,ta tong shi ju you xian zhe de peng zhang jiao yuan dan bai de neng li ,zhe biao ming myroilysinzai sheng wu ji shu ying yong fang mian ju you hen da qian li 。you yu myroilysinbu neng zi ti cheng shou ,yin ci di gao ye sheng M.profundi D25xi jun zhong myroilysinde chan liang you ji chong yao 。zai ben lun wen zhong ,wo men shou xian li yong chan yin zi shi yan you hua le jun zhu Mrofundi D25de pei yang tiao jian 。sui hou ,li yong you hua hou de pei yang tiao jian ,wo men jin hang le duo ci jun zhu M.profundi D25de xiao shi he zhong shi fa jiao ,bing cheng gong jian li le M.profundi D25chan myroilysinde xiao shi he zhong shi fa jiao gong yi 。fa jiao gong yi de jian li ,wei dan bai mei myroilysinde gong ye hua sheng chan he ji sheng wu ji shu fang mian de kai fa ying yong dian ding le ji chu 。
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