專(zhuān)利名稱(chēng)：：人源5,10-次甲基四氫葉酸合成酶及其復(fù)合物的結(jié)晶方法、晶體以及應(yīng)用的制作方法
技術(shù)領(lǐng)域：
：本發(fā)明涉及一種人源5，10-次甲基四氫葉酸合成酶(MTHFS)的晶體及其結(jié)晶的方法，同時(shí)也涉及人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與產(chǎn)物10-甲酰四氫葉酸復(fù)合物的晶體及其結(jié)晶的方法；還涉及這些晶體中對(duì)于人源5，10-次甲基四氫葉酸合成酶的酶活性起關(guān)鍵作用的結(jié)構(gòu)域或肽片段；以及利用這些重要結(jié)構(gòu)域或肽片段進(jìn)行藥物設(shè)計(jì)和篩選的方法。
背景技術(shù)：
：葉酸，這種水溶性的B族維生素早在19世紀(jì)40年代已被發(fā)現(xiàn)。葉酸介導(dǎo)的單碳代謝途徑存在于從微生物到高等植物、動(dòng)物的生命活動(dòng)中，此代謝途徑對(duì)細(xì)胞的發(fā)育和增殖非常重要，并且在很多生理活動(dòng)中起著非常重要的作用，因此還原型葉酸是其中重要的輔助因子。葉酸只能在原核生物、酵母、植物中合成，但是對(duì)人體卻是必須的，葉酸介導(dǎo)的單碳代謝途徑如果被阻斷，在人體內(nèi)將引發(fā)癌癥，心血管疾病和發(fā)育異常等。該代謝途徑中涉及的酶類(lèi)，很久以來(lái)就被用作治療癌癥(A卯ling，1991;FoxandStrover，2008;Chattopadhyayet.al.，2007;Farberet.al.，1948)。目前，抗葉酸藥物不僅被用作化療藥物，也在治療風(fēng)濕性關(guān)節(jié)炎和銀屑病等疾病中發(fā)揮作用。葉酸在生物體內(nèi)以多種氧化態(tài)存在，如葉酸，二氫葉酸，四氫葉酸等。這些不同狀態(tài)的葉酸攜帶的單碳單位也存在不同的氧化狀態(tài)如甲醇或甲酸，它們通過(guò)酶學(xué)反應(yīng)進(jìn)行相互的轉(zhuǎn)換。不同狀態(tài)的四氫葉酸可以接受或提供單碳單位給多種生命活動(dòng)。包括(l)嘌呤和嘧啶的從頭合成，二磷酸尿嘧啶甲基化形成二磷酸胸腺嘧啶；(2)同型半胱氨酸甲基化形成甲硫氨酸，甲硫氨酸可以形成s-腺苷甲硫氨酸，后者對(duì)細(xì)胞內(nèi)的甲基化反應(yīng)至關(guān)重要。嘌呤、嘧啶的從頭合成和同型半胱氨酸的合成都對(duì)代謝途徑上游的合成阻斷很敏感，會(huì)導(dǎo)致二磷酸尿嘧啶錯(cuò)誤地?fù)饺隓NA當(dāng)中，并且會(huì)提升細(xì)胞內(nèi)的同型半胱氨酸濃度，破壞s-腺苷甲硫氨酸依賴型甲基化反應(yīng)。葉酸代謝途徑的紊亂將導(dǎo)致多重嚴(yán)重疾病，所以該途徑的很多酶都被作為藥物設(shè)計(jì)的重要靶點(diǎn)。葉酸的單谷氨酸衍生物穿過(guò)細(xì)胞膜進(jìn)入細(xì)胞，在細(xì)胞內(nèi)被多聚谷氨酸合成酶催化合成不同長(zhǎng)度的谷氨酸衍生物，多聚谷氨酸支鏈可以使葉酸存在于細(xì)胞內(nèi)，同時(shí)也可以增加其與胞內(nèi)結(jié)合酶類(lèi)的結(jié)合能力。5，10-次甲基四氫葉酸合成酶(Homosapiens5，lO-methenyltetrahydrofolatesynthetase,簡(jiǎn)稱(chēng)MTHFS)，也稱(chēng)為5_甲酰四氫葉酸環(huán)化酶(5-formyltetrahydrofolatecyclo-ligase)。酶學(xué)分類(lèi)號(hào)(EC)6.3.3.2.MTHFS催化的體內(nèi)代謝反應(yīng)為55-甲酰四氫葉酸5,10-次甲基四氫葉酸反應(yīng)的底物為5-甲酰四氫葉酸(5-formyltetrahydrofolate，簡(jiǎn)稱(chēng)5-FTHF)，反應(yīng)的產(chǎn)物為5，10-次甲基四氫葉酸(5，10-methenyltetrahydrofolate，簡(jiǎn)稱(chēng)5，10-MTHF)。此催化反應(yīng)在室溫下即可進(jìn)行，二價(jià)鎂離子和ATP(三磷酸腺苷)為5，10-次甲基四氫葉酸合成酶的輔助因子。MTHFS存在于很多生物中，包括細(xì)菌、植物和動(dòng)物，表明其在葉酸代謝中具有很重要的作用。目前對(duì)葉酸代謝途徑中多種重要蛋白的酶學(xué)特征研究結(jié)果顯示，5_甲酰四氫葉酸是目前已知的唯一熱動(dòng)力學(xué)穩(wěn)定的還原型葉酸衍生物，并且沒(méi)有被直接用作單碳載體。在原核生物中，5_甲酰四氫葉酸被認(rèn)為是一種單碳載體的儲(chǔ)存狀態(tài)，主要存在于種子和孢子中，其狀態(tài)隨5，10-次甲基四氫葉酸合成酶的表達(dá)而變化。在哺乳動(dòng)物中，5-甲酰四氫葉酸占胞內(nèi)總?cè)~酸含量的10%-15%，5-甲酰四氫葉酸對(duì)一些單碳代謝途徑中的酶類(lèi)有抑制作用，如絲氨酸羥甲基轉(zhuǎn)移酶(serinehydroxymethyltransferase)和5-氨基咪唑_4_甲酰胺基核苷酸轉(zhuǎn)甲?；?AICARTfase)。5，10-次甲基四氫葉酸合成酶是唯一可以將5-甲酰四氫葉酸轉(zhuǎn)換為其它類(lèi)型還原型葉酸的酶。通過(guò)調(diào)節(jié)體內(nèi)5-甲酰四氫葉酸的濃度，5，10-次甲基四氫葉酸合成酶可以控制很多葉酸依賴型代謝途徑，例如嘌呤、嘧啶、氨基酸等代謝途徑。上述代謝途徑與生命發(fā)育，癌癥預(yù)防和治療中的關(guān)鍵步驟緊密相連。有研究表明使用5-甲酰四氫同型葉酸，可以抑制5，10-次甲基四氫葉酸合成酶的活性，進(jìn)而抑制人乳腺癌。從圖1中可以看到，5，10-次甲基四氫葉酸合成酶在單碳代謝途徑中的作用，它同時(shí)可以正向和反向調(diào)節(jié)此代謝途徑。5，10-次甲基四氫葉酸合成酶是單碳循環(huán)上游的關(guān)鍵酶，通過(guò)代謝途徑的分析，它對(duì)下游代謝的影響要比二氫葉酸還原酶顯著。對(duì)于癌癥引起的基因組甲基化異常以及DNA及RNA的完整性、DNA的修復(fù)能力等方面來(lái)說(shuō)，5，10-次甲基四氫葉酸合成酶都是一個(gè)很關(guān)鍵的調(diào)控靶蛋白。5，10-次甲基四氫葉酸合成酶在不同生物中的氨基酸序列與人體內(nèi)的氨基酸序列同源性比較列于表1中(同源性比較使用NCBI的BLAST軟件)表l<table>tableseeoriginaldocumentpage7</column></row><table>雖然原核生物中的5，10-次甲基四氫葉酸合成酶三維結(jié)構(gòu)已經(jīng)被解析，其與人源5，10-次甲基四氫葉酸合成酶氨基酸序列的同源性僅為28%。而哺乳動(dòng)物以及人的5，10-次甲基四氫葉酸合成酶的三維結(jié)構(gòu)解析卻未見(jiàn)報(bào)道。由于葉酸的衍生物在生理環(huán)境中不穩(wěn)定，體內(nèi)多數(shù)葉酸衍生物是以結(jié)合在蛋白上的形式存在的。葉酸依賴型單碳代謝網(wǎng)絡(luò)中酶反應(yīng)鏈中存在著底物流傳遞，即底物與酶結(jié)合后發(fā)生反應(yīng)，反應(yīng)完成后的產(chǎn)物并不與酶分離，而是以這種結(jié)合狀態(tài)傳遞到下一個(gè)酶反應(yīng)中。研究也表明，這一代謝途徑中通過(guò)形成中間態(tài)來(lái)調(diào)節(jié)酶活性，進(jìn)而調(diào)節(jié)代謝途徑的趨向和保持動(dòng)態(tài)平衡。如從豬肝中提取的10-甲酰四氫葉酸脫氫酶，就是已結(jié)合有帶6個(gè)谷氨酸的四氫葉酸。這樣就導(dǎo)致如果反應(yīng)鏈中一個(gè)酶發(fā)生變異，其中間產(chǎn)物發(fā)生積累，則對(duì)整個(gè)代謝途徑的影響都是深遠(yuǎn)的。MTHFS催化5-甲酰四氫葉酸不可逆轉(zhuǎn)化為5，10-次甲基四氫葉酸(Jolivet，1997)，此催化反應(yīng)依賴于三磷酸腺苷(adenosine-triphosphate,簡(jiǎn)稱(chēng)ATP)并且可被產(chǎn)物反饋抑制(Huennekenset.al.，1984;Jolivetet.al.，1996;Maraset.al.，1994)。核磁共振(NMR)波譜和同位素交換方法研究表明MTHFS催化的反應(yīng)存在兩個(gè)中間態(tài)(HuangandSchirch，1995)。(1)5-甲酰四氫葉酸的甲?；鶊F(tuán)親核攻擊ATP的磷酸基團(tuán)產(chǎn)生第一個(gè)N5-亞胺磷酸中間態(tài)，這部分反應(yīng)是可逆的。(2)第二個(gè)親核攻擊發(fā)生在分子內(nèi)，N10與N5-亞胺磷酸的N5連接，形成一個(gè)四面體中間態(tài)磷酸咪唑啉。磷酸咪唑啉中間態(tài)移除磷酸基團(tuán)，最后形成催化產(chǎn)物5，10-次甲基四氫葉酸。5，10-次甲基四氫葉酸是葉酸代謝的中間產(chǎn)物，可以在與酶結(jié)合的狀態(tài)下轉(zhuǎn)化為10-甲酰四氫葉酸。反應(yīng)的最后一步使得由MTHFS催化的反應(yīng)不可逆。目前，對(duì)這一反應(yīng)機(jī)理的三維結(jié)構(gòu)方面的信息所知甚少。有報(bào)道的是源于肺炎支原體(Mycoplasm即皿emoniae)的MTHFS(mpMTHFS)，并且結(jié)合二磷酸腺苷(adenonisinedisphosphate，簡(jiǎn)稱(chēng)ADP)和5-甲酰四氫葉酸的三元復(fù)合物結(jié)構(gòu)(Chenet.al.，2005)，另外一個(gè)是源于炭疽芽胞桿菌(Bacillusanthracis)(bMTHFS)的MTHFS，并且結(jié)合有ADP(Meieret.al.，2007)。人源MTHFS(hMTHFS)的氨基酸序列與蛋白質(zhì)數(shù)據(jù)庫(kù)(http:〃www.rcsb.org/pdb/)中的一級(jí)結(jié)構(gòu)相比較同源性少于28%(Bermanet.al.，2000)。從已知的結(jié)構(gòu)看出hMTHFS和mpMTHFS、bMTHFS的整體結(jié)構(gòu)十分相似，預(yù)示著MTHFS趨向于一種保守的折疊方式。在鼠類(lèi)研究中，利用酶穩(wěn)態(tài)動(dòng)力學(xué)和分子模擬技術(shù)表明，N5和N10甲基化的葉酸衍生物用做MTHFS抑制物的可行性較小(Fieldet.al.，2006,2007)。在直腸癌和胰癌化療中常規(guī)使用的5-氟尿嘧啶(5-fluorouracil，簡(jiǎn)稱(chēng)5-FU)是通過(guò)抑制胸苷酸合成酶(thymidylatesynthase,簡(jiǎn)稱(chēng)TS)發(fā)揮作用的(Longleyet.al.，2003)，胸苷酸合成酶(TS)從5，10-亞甲基四氫葉酸上獲得一個(gè)單碳單元來(lái)催化脫氧尿苷酸(deoxyuridylate)轉(zhuǎn)化為脫氧胸苷酸(deoxythymidylate)，這一反應(yīng)對(duì)于DNA的合成至關(guān)重要(CarrerasandSanti，1995)。臨床上，為了提高治療效果，5-甲酰四氫葉酸(5-formylTHF)通常和5_FU—起使用。這樣使用的好處在于MTHFS可以催化5-甲酰四氫葉酸合成5，10-次甲基四氫葉酸，而5，10-次甲基四氫葉酸可以轉(zhuǎn)化為5，10-亞甲基四氫葉酸。低含量的5，10-亞甲基四氫葉酸導(dǎo)致5-FU從TS四聚體上解離下來(lái)，從而降低胸苷類(lèi)似物的抑制效果(Bertino，1977;Sirotnaket.al.，2000)。5，10-亞甲基四氫葉酸也可由絲氨酸羥甲基轉(zhuǎn)移酶(serinehydroxymethyltransferase，簡(jiǎn)稱(chēng)SHMT)在轉(zhuǎn)化絲氨酸為甘氨酸的過(guò)程中合成。SHMT的酶活性可以被MTHFS直接調(diào)節(jié)，因?yàn)镸THFS的底物5-甲酰四氫葉酸可以抑制SHMT(Renwicket.al.，1998;SchirchandSzebenyi，2005;StoverandSchirch，1990)。5-氨基咪唑-4-甲酰胺基核苷酸轉(zhuǎn)甲?；?phosphoribosylaminoimidazolecarboxamidetransformylase,簡(jiǎn)稱(chēng)AICARFT)和甘氨酰胺核苷酸轉(zhuǎn)甲?；?Glycinamideribonucleotidetransformylase,簡(jiǎn)稱(chēng)GARFT)是單碳代謝途徑中參與嘌呤代謝的兩個(gè)酶，可以從10-甲酰四氫葉酸上獲得單碳單元，也被用來(lái)作為抗癌藥物的靶點(diǎn)(Allegraet.al.，1985;Piperet.al.，1988)。洛美曲索(Lometrexol)為第二代葉酸拮抗劑的代表，用來(lái)抑制GARFT活性，進(jìn)而抑制嘌呤合成，達(dá)到治療癌癥的目地。使用甲氨蝶呤(Methotrexate,簡(jiǎn)稱(chēng)MTX)和氨基蝶呤(aminopterin)抑制二氫葉酸還原酶(dihydrofolatereductase，簡(jiǎn)稱(chēng)DHFR)導(dǎo)致二氫葉酸的多聚谷氨酸衍生物積累，進(jìn)而抑制AICARFT的酶活性(Quinlivanet.al.，2000;Shihet.al.，1997)。同樣，通過(guò)5-甲酰四氫葉酸的類(lèi)似物抑制5，10-次甲基四氫葉酸合成酶(5，10-methenyltetrahydrofolatesynthetase,簡(jiǎn)稱(chēng)MTHFS)導(dǎo)致5_甲酰四氫葉酸的積累，進(jìn)而抑制絲氨酸羥甲基轉(zhuǎn)移酶(serinehydroxymethyltransferase,簡(jiǎn)稱(chēng)SHMT)和AICARFT的酶活性。如上所述，酶活性的抑制對(duì)碳單元在代謝途徑中的轉(zhuǎn)移是有負(fù)面效應(yīng)的，導(dǎo)致細(xì)胞中一些重要組分的合成阻斷，這也是目前使用葉酸類(lèi)似物進(jìn)行癌癥化學(xué)治療的機(jī)理。使用葉酸衍生物進(jìn)行癌癥治療，存在兩個(gè)問(wèn)題，(一)這些藥物經(jīng)過(guò)幾十年的使用后，在很多病例中發(fā)現(xiàn)了耐藥現(xiàn)象，而且對(duì)一些病人來(lái)說(shuō)，這兩種藥物效果不夠明顯；(二)這些藥物多具有比較強(qiáng)的細(xì)胞毒性。因此，開(kāi)發(fā)新型葉酸衍生物藥物用于癌癥化學(xué)治療是目前的研究熱點(diǎn)。
發(fā)明內(nèi)容本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶的晶體，其特征在于所述晶體的空間群為C222l，晶胞參數(shù)為a=48.70±2A，b=145.12±2A，c=59.21±2A;a=90.oo°，e=90.oo°，y=90.oo°。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a=48.51±2A,b=145.22±2人，c=59.65±2人；a=90.00°，e=90.00°，y=90.00°。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，其特征在于所述晶體的空間群為P21212，晶胞參數(shù)為<formula>formulaseeoriginaldocumentpage9</formula>。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為<formula>formulaseeoriginaldocumentpage9</formula>本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為<formula>formulaseeoriginaldocumentpage9</formula>本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為<formula>formulaseeoriginaldocumentpage9</formula>本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶的晶體的結(jié)晶方法，包括將所述人源5，10-次甲基四氫葉酸合成酶的溶液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在l-2天后收集晶體；其特征在于在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP溶液，再加入還原劑和鎂離子，混合后得到人源5，10-次甲基四氫葉酸合成酶與ATP的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與ATP的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入谷氨酸溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與谷氨酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與谷氨酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP、5_甲酰四氫葉酸、巰基乙醇、鎂離子溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪_N_2-乙磺酸或嗎啉乙磺酸緩沖液。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP、5_甲酰四氫葉酸、巰基乙醇、鎂離子溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在4-14天后收集晶體；其特征在于在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含O.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪_N_2-乙磺酸或嗎啉乙磺酸緩沖液。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶的晶體的F55、M58、E61、M90、￥152、￥153位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與底物5-甲酰四氫葉酸中的蝶呤基團(tuán)結(jié)合。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶的晶體的F85、W109、R148位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與底物5-甲酰四氫葉酸中的對(duì)氨基苯甲酸基團(tuán)結(jié)合。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶的晶體的K10、R14、R145、D154位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與ATP結(jié)合。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體中a5的152-161位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域是與從ATP水解下來(lái)的磷酸基團(tuán)結(jié)合的結(jié)構(gòu)域。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體中P3和P4之間的氨基酸即第89和135位氨基酸之間的氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域形成一個(gè)loop環(huán)結(jié)構(gòu)，該結(jié)構(gòu)中的氨基酸與谷氨酸結(jié)合。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體中氨基酸殘基F55，L56，S57，M58，E61，P81，Y83，M90，W109，P135，R148，K150，Y152和Y153所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域形成一個(gè)球形口袋，N5-亞胺磷酸過(guò)渡態(tài)的蝶呤環(huán)位于所述球形口袋的中間。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體中蝶呤環(huán)在空間中的結(jié)構(gòu)域，其特征在于所述蝶呤環(huán)以C4a與C8a之間的鍵為軸發(fā)生了扭轉(zhuǎn)，所述喋呤環(huán)上的電子發(fā)生了重排。本發(fā)明提供一種人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體中Y83在空間中的結(jié)構(gòu)域，其特征在于Y83的位置與在權(quán)利要求5所述的晶體的位置相比，發(fā)生了變化。本發(fā)明提供與上述的任一結(jié)構(gòu)域類(lèi)似或者結(jié)合的小分子化合物、短肽、多肽、核酸、抗體或免疫結(jié)合物中的一種或幾種的組合物。圖1示出了5，10-亞甲基四氫葉酸合成酶在單碳代謝途徑中的位置及作用。TS:胸苷酸合成酶(thymidylatesynthase,簡(jiǎn)稱(chēng)TS);MTHFS:5，10-次甲基四氫葉酸合成酶(5，10-methenyltetrahydrofolatesynthetase，簡(jiǎn)稱(chēng)MTHFS);SHMT:絲氨酸羥甲基轉(zhuǎn)移酶(serinehydroxymethyrltransferase，簡(jiǎn)稱(chēng)SHMT);DHFR:二氫口十酸還原酶(dihydrofolatereductase,簡(jiǎn)稱(chēng)DHFR);MTHFR:亞甲基四氫葉酸還原酶(Methylenetetrahydrofolatereductase,簡(jiǎn)稱(chēng)MTHFR);MTHFD:亞甲基四氫葉酸脫氫酶(methylenetetrahydrofolatedehydrogenase,簡(jiǎn)稱(chēng)MTHFD)圖2示出了MTHFS催化反應(yīng)產(chǎn)物5，10-次甲基四氫葉酸轉(zhuǎn)化為10-甲酰四氫葉酸，5，10-次甲基四氫葉酸是葉酸代謝的中間產(chǎn)物，可以在與酶結(jié)合狀態(tài)轉(zhuǎn)化為10-甲酰四氫葉酸。A:5，10-次甲基四氫葉酸；B:10-甲酰四氫葉酸。圖3示出了5，10-亞甲基四氫葉酸合成酶晶體的立體三維結(jié)構(gòu)示意圖。圖4示出了5，10-亞甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體的三維結(jié)構(gòu)示意圖。圖5示出了5，10-亞甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體的三維結(jié)構(gòu)示意圖。圖6示出了5，10-亞甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體的三維結(jié)構(gòu)示意圖。圖7示出了5，10-亞甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體的三維結(jié)構(gòu)示意圖。具體實(shí)施例方式人源5，10-次甲基四氫葉酸合成酶(MTHFS)的表達(dá)與純化MTHFS的氨基酸序列如下MAAAAVSSAKRSLRGELKQRLRAMSAEERLRQSRVLSQKVIAHSEYSTA從商品化人轉(zhuǎn)錄文庫(kù)中PCR克隆得到全長(zhǎng)的人MTHFS的基因。使用的引物如下BCO19921上游TACTTCCAATCCAATGCTATGGCGGCGGCAGCGGTBC019921下游TTATCCACTTCCAATGTTAAGCTGTTGACGAGTCT克隆得到的基因，通過(guò)連接反應(yīng)克隆到商品化PET2lb載體中，該載體在蛋白質(zhì)的氮端具有6個(gè)組氨酸的表達(dá)標(biāo)簽。使用BL21(DE3)宿主菌表達(dá)重組蛋白，37攝氏度，當(dāng)菌濃度達(dá)到0D6。Qnm=0.8時(shí)，加入終濃度為0.2mM的IPTG。然后，16攝氏度表達(dá)24小時(shí)，4000rpm，收集表達(dá)菌。收集的表達(dá)菌重懸于磷酸鹽緩沖液(50mMNa2HP04，10mMKH2P04，137mMNaCl，2.7mMKC1，pH7.4)，使用100W超聲波破碎細(xì)胞，在磷酸鹽緩沖液中重懸細(xì)菌，16000rpm，30分鐘，取上清，經(jīng)QIAGEN的Ni-NTA親和層析柱可以得到初步純化的蛋白，使用煙草蝕紋病毒(TEV)蛋白酶切除酶氮端具有的6個(gè)組氨酸表達(dá)標(biāo)簽。然后，再通過(guò)QIAGEN的Ni-NTA親和層析柱去除蛋白中的TEV蛋白酶及6個(gè)組氨酸表達(dá)標(biāo)簽。獲得的純化蛋白，使用Milipore公司的濃縮產(chǎn)品Amicon、截留分子量為5kDa，濃縮蛋白到1毫升，使用GE公司的SuperDexG75分子篩純化。使用分子篩純化MTHFS為單一的洗出峰，同時(shí)蛋白緩沖液換為20mMTris-HCl，pH8.0，200mMNaCl，0.5mMEDTA(乙二胺四乙酸二鈉)和5mMP-巰基乙醇-merc即toethanol)，使用Milipore公司的濃縮產(chǎn)品Amicon，截留分子量為5kDa，濃縮至lj10mg/ml。晶體的生長(zhǎng)晶體可采用分批結(jié)晶、液-液擴(kuò)散、蒸氣擴(kuò)散、透析等方法獲得。本發(fā)明中采用的是液_液擴(kuò)散的方法，包括懸滴法或坐滴法。在以下的實(shí)驗(yàn)中采用Hampton公司的晶體篩選試劑盒，晶體生長(zhǎng)的條件為在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH=6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。在結(jié)晶前，將人源5，10-次甲基四氫葉酸合成酶與池液混合或者將人源5，10-次甲基四氫葉酸合成酶和各種底物的混合液與池液混合。下面披露一些優(yōu)選的實(shí)施方式人源5，10-次甲基四氫葉酸合成酶的晶體的結(jié)晶方法，首先將所述人源5，10-次甲基四氫葉酸合成酶的濃縮液與池液混合；將上述酶蛋白溶液在Eppendorf冷凍離心機(jī)離心14，OOOrprn，15分鐘；采用Hampton公司的晶體篩選試劑盒結(jié)晶，結(jié)晶的池液的組成為0.02MMgCl2、0.02MNiCl2、HEPES(O.1M，p朋.6)和20%PEG3350，大約兩天后收集得到的晶體。人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體的結(jié)晶方法，首先向所述人源5，10-次甲基四氫葉酸合成酶的溶液加入終濃度為5mM的ATP溶液，再加入還原劑和鎂離子，混合后得到人源5，10-次甲基四氫葉酸合成酶與ATP的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與ATP的混合液與池液混合1小時(shí)；結(jié)晶前將所述蛋白溶液在E卯endorf冷凍離心機(jī)離心14，OOOrprn，15分鐘(主要目的是為了去除雜質(zhì))，采用Hampton公司的晶體篩選試劑盒結(jié)晶，結(jié)晶的池液的組成為O.02MMgCl2、0.02MNiCl2、HEPES(0.1M，p朋.6)和20%PEG3350,1_2天收集晶體進(jìn)行數(shù)據(jù)收集。人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體的結(jié)晶方法，向5-10mg/ml蛋白溶液中加入終濃度為3mM的谷氨酸溶液，混合1小時(shí)，結(jié)晶前將所述蛋白溶液在E卯endorf冷凍離心機(jī)離心14，OOOrpm，15分鐘，采用Hampton公司的晶體篩選試劑盒結(jié)晶，結(jié)晶的池液的組成為0.02MMgCl2、0.02MNiCl2、HEPES(O.1M，p朋.6)和20%PEG3350,1-2天收集晶體進(jìn)行數(shù)據(jù)收集。人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體的結(jié)晶方法，向5-10mg/ml蛋白溶液中加入終濃度為5mMATP、5mM5-甲酰四氫葉酸、2mM巰基乙醇、5mM鎂離子，混合1小時(shí)，結(jié)晶前將所述蛋白溶液在Eppendorf冷凍離心機(jī)離心14，000rpm，15分鐘，采用Hampton公司的晶體篩選試劑盒結(jié)晶，結(jié)晶的池液的組成為0.02MMgCl2、0.02MNiCl2、HEPES(0.1M，p朋.6)和20%PEG3350,1-2天收集晶體進(jìn)行數(shù)據(jù)收集。人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體的結(jié)晶方法，5-10mg/ml蛋白溶液中加入終濃度為5mMATP、5mM5-甲酰四氫葉酸、2mM巰基乙醇、5mM鎂離子，混合1小時(shí)，結(jié)晶前將所述蛋白溶液在E卯endorf冷凍離心機(jī)離心14，OOOrpm，15分鐘，結(jié)晶的池液的組成為0.02MMgCl2、0.02MNiCl2、HEPES(O.1M，p朋.6)和20%PEG3350，4-14天收集晶體進(jìn)行數(shù)據(jù)收集。結(jié)晶的具體過(guò)程為晶體生長(zhǎng)于16t:，進(jìn)行懸滴蒸氣擴(kuò)散結(jié)晶或坐滴法結(jié)晶實(shí)驗(yàn)，將包含混合池液2微升和2微升蛋白溶液或蛋白溶液與底物或產(chǎn)物的混合液的混合，進(jìn)行擴(kuò)散，在300iil包含O.02MMgCl2、0.02MNiCl2、HEPES(0.lM，p朋.6)和20%PEG3350的池液存在下，經(jīng)過(guò)一段時(shí)間發(fā)現(xiàn)最大尺寸達(dá)300-500i！m的片狀晶體，將晶體轉(zhuǎn)移至包含0.01MMgC12、0.01MNiC12、HEPES(0.lM，p朋.6)、10%PEG3350、20X甘油作為防凍劑的溶液中。然后在收集X-射線數(shù)據(jù)前快速冷凍樣品。晶體的解析獲得的復(fù)合物晶體轉(zhuǎn)移到含有10_15%甘油的池液中，挑出單晶，快速冷凍至液氮中，經(jīng)初步X衍射篩選后，確定要收集的晶體數(shù)據(jù)。衍射數(shù)據(jù)的收集使用同步輻射，波長(zhǎng)為0.979A，數(shù)據(jù)初步處理使用HKL2000(0twinowskiandMinor,1997)。野生型結(jié)構(gòu)解析使用分子置換方法，使用軟件為Balbes(Longet.al.，2008)，以蛋白質(zhì)數(shù)據(jù)庫(kù)中的mpMTHFS(PDBcode1U3G)建模。復(fù)合物晶體數(shù)據(jù)收集使用in-houseFRE+copperrotatinganodeandaR-AXISIV++detector(Rigaku，USA)及同步輻射。復(fù)合物結(jié)構(gòu)的解析使用軟件為Phaser(McCoyet.al.，2007)，以人源hMTHFS的結(jié)構(gòu)為模板建模。結(jié)構(gòu)模型的修正使用軟件Phenixrefine(Adamset.al.，2002)和Refmac(Murshudovet.al.，1997)。用R因子和Rfree因子來(lái)監(jiān)測(cè)模型質(zhì)量和修正誤差。人源5，10-次甲基四氫葉酸合成酶的晶體，空間群為C2221，晶胞參數(shù)為a=48.70±2A,b=145.12±2A，c=59.21±2A;a=90.00°，P=90.00°，y=90.00°。人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，空間群為C2221，晶胞參數(shù)為a=48.51±2A，b=145.22±2A,c=59.65±2A;a=90.00°，e=90.00°，y=90.00°。人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，空間群為P21212，晶胞參數(shù)為a=59.579±2A,b=144.101±2A,c=48.532±2A;a=90.00°，P=90.00°，y=90.00°。人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體，空間群為C2221，晶胞參數(shù)為a=48.45±2A,b=145.23±2A,c=59.60±2A;a=90.00°，e=90.00°，y=90.00°。人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體，空間群為C222l，晶胞參數(shù)為a=48.69±2A,b=141.03±2A,c=59.99±2A;a=90.00°，P=90.00°，y=90.00°。人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體，空間群為C222l，晶胞參數(shù)為a=48.86土2A,b=144.97±2A,c=59.41±2A;a=90.00°，P=90.00°，y=90.00°。表2為人源5，10-次甲基四氫葉酸合成酶的晶體的坐標(biāo)表2<table>tableseeoriginaldocumentpage14</column></row><table>48NALAA935.63352.207-7.26210029.5949CAALAA935.24451.141-6.34910019.0350CBALAA935.76851.432—4.95510027.2851CALAA933.72751.002-6.33410023.14520ALAA933.19449.897-6.42810029.0753NLYSA1033.03952.134-6.23610022.0754CALYSA1031.58052.146-6.21210025.8855CBLYSA1031.05153.566-5.97010032.8656CGLYSA1031.32954.113_4.57310022.9957CDLYSA1030.75155.518-4.39910021.0258CELYSA1030.61655.880-2.92510032.3659NZLYSA1029.96657.207-2.72610043.9660CLYSA1030.99751.578-7.50310025.13610LYSA1030.08550.750-7.47410025.5662NARGA1131.53152.024-8.6361.0023.0963CAARGA1131.06551.553-9.9301.0020.]764CBARGA1131.81752.271-ll.0511.0033.5065CGARGA1131.44651.815-12.4461.0032.4366CDARGA1132.30652.533-13.4741.0037.1267NEARGA1133.71252.514-13.0801.0035.8368CZARGA1134.51651.468-13.2401.0046.9269NH1ARGA1134.05550.354-13.7941.0047.6970NH2ARGA1135.78251.534-12.8481.0043.7971CARGA1131.22950.045-IO.0731.0021.69720ARGA1130.325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子15430HOH19.58367.0671.0991004.200原子15440HOH29.96270.0550.51110016.460原子15450HOH38.03057.515-6.47310019.940原子15460HOH411.74656.5610.95910015.980原子15470HOH50.92854.112-1147610017.610原子15480HOH69.52755.580-525710019.760原子15490■79.17472.967-2.61810020.160原子15500匪8-O.14262.909-10.97110019.760原子15510HOH90.83154.30913.61210023.810原子15520HOH1011.04671.372-2.86210013.360原子15530HOH1116.15646.268-l.68810021.530原子15540HOH1223.34157.589-5.00710027.920原子15550國(guó)1311.29442.957一640310032.980原子15560H0H140.00054.788-14.8041.0029.190原子15570HOH153.50666.394-8.2401.0024.860原子15580HOH166,01249.126-4.1181.0026.620原子15590HOH1710.08666.627-7.2991.0023.300原子15600HOH1813.64272.4632.34610031.110原子15610HOH190.36266.4017.18010025.870原子15620■204.皿70.052-4.88010024.350原子15630■21-5.03755.9227.05010034.820原子15640■2212.23652.918-5.74610029.850原子15650HOH2318.67159.07012.36910043.000原子15660HOH241.31546.97716.5781.0035.890原子15670HOH25-3.19757.7666.9771.0029.530原子15680■2610.38969.271-7.1571.0017.050原子15690HOH2725.52757.1340.0971.0035.690原子15700HOH280.05945.27511.3521.0028.910原子15710■2911.37657.034-IO.6541.0022.350原子15720HOH306.40545.0941.5801.0026.760原子15730■31-3.83162.752-2.9311.0032.780原子15740■3211.14160.326-16.0701.0026.740原子15750HOH336.73572.5610.0000.7711.810原子15760HOH349.53058.150-8.8051.0022.020原子15770HOH3516.42739.8862.9761.0028.660原子15780HOH360.13848.701-1.2621.0027.300原子15790匪371.46451.70214.6031.0024.790原子15800HOH3831.31242.328-O.5521.0032.800原子15810HOH3912.36446.39717.0671.0035.330原子15820HOH403.70849.731-4.7601.0026.440原子15830HOH413.53371.268-0.3361.0027.240原子15840麗426.76364.585-11.5571.0030.920原子15850HOH43-8.41956.456-2.3701.0029.900原子15860HOH442.07472.3362.7231.0031.620原子15870HOH4513.20630.466-7.2491.0033.580原子15880HOH467.43967.126-11.0721.0036.310原子15890HOH475.49761.144-13.2581.0030.700原子15900■48-0.86344.6147.4041.0031.320原子15910HOH496.41749.742-8.2871.0033.540原子15920HOH5016.43659.82914.9531.0035.850原子15930■519.81266.574-10.1761.0027.700原子15940國(guó)52-7.88064.6261.8741.0056.140原子15950■5316.30948.928_1.7401.0031.700原子15960HOH5432.31044.685—0.23310036.380原子15970■557.59064.45912.11210032.700原子15980■567.91070.302_8.30610034.010原子15990HOH5718.21440.463-9.56310028.630原子16000■5819.49339.98910.35110045.090原子16010HOH5911.71043.47713.55910031.770原子16020HOH6012.29955.406-6.24110032.820原子16030HOH61-6.74166.7160.52810042.370原子16040國(guó)628.50070.7118.24710026.350原子16050■636.67059.62117.45510048.640原子16060HOH6431.04356.Oil-9.09610039.000原子16070HOH6520.16058.4589.25510044.830原子16080■669.81864.20413.6520044.890原子16090HOH67-1.06259.8313.96910020.690原子16100■6818.89338.320-11.92510035.040原子16110HOH6929.34358.240-O.09710042.870原子16120■7017.10665.358_3.01510024.460原子16130HOH7110.46570.738-15.25710043.180表3為人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體的坐標(biāo):表316140H0H7229.56958.034-7.5051.0039..6516150HOH73-3.44859.6844.9981.0034.6216160HOH745.54743.502-0.6211.0037.0616170HOH751.03249.483-3.6531.0040.5116180HOH764.88859.84820,5261.0035.5716190HOH7718.00742.0469.9191.0036.2216200HOH7833.48060.828-10.0121.0044.1516210HOH7927.71960.1961.6171.0038.5116220HOH8027.60041.102-2.1951.0041.8916230■8122.00647.62910.8671.0034.2716240HOH8218.46633.821-2.1191.0038.1216250HOH8320.98636.410-10.9281.0041.1316260HOH8414.09530.178-1.7621.0038.0216270HOH85-2.57867.1241.8611.0037.0316280■8619.97858.288_3.9841.0020.0616290關(guān)87-6.57654.9272.6011.0039.5716300腳8817.17230.892-O.5141.0043.6816310HOH89_6.78163.9604.2841.0045.1816320HOH905.59041.3301.1441.0036.0416330■9110.58869.78213.6441.0056.5616340HOH9234.93847.666-5.2521.0038.6316350HOH9328.13553.804-1.5721.0042.146360關(guān)9423.82349.73711.3921.0054.1616370HOH9510.32039.781-11.2631.0049.6916380HOH9619.88443.016-7.1881.0052.4616390HOH9711.17426.833-8.0581.0049.8616400■9816.77057.623一9.6081.0042.9416410HOH99-2.27266.7328.1721.0059.6916420HOH10019.43544.20810.8741.0038.3216430關(guān)10117.53330.4192.4701.0047.4816440HOH1024,39941.716—2.6471.0042.210000000000000000000000000000000子子子子0035.43N0041.41C0045.63C0045.32c0037.59c0059.03s0053.80c0042.0900049.5300048.40N0045.74c0049.29c0040.17c0042.9000048.58N0052.51c0050.38c0047,96c0036.70N0040.63c0032.11c0039.83c0035.2300029.09N0031.34c0040.09c0041.10c0035.6800038.33N0048.93c0060.52c0047.51c0034.39c0031.98c0028.6800042.71N0034.73c0035.76c0051.4600040.44c0039,3500034.96N0035.73c0030.41c0041.6800040.24c0031.0400027.74N0028.91c0036.07c0030.07c0036.1700032.46N0040.86c0041.64c0038.00c0034.36c0032.98cCN101781637A說(shuō)明書(shū)A1-17348-10.1837.583A1-17.905-11.5157.788A1-17380-12.4766.724A1-19.440-11.4787.773A1一20.067-10.6178.868A1-19.725-11.22810.533A1-20.654-12.75710.528A1-16.773-13.4997.044A2-14.960-14.0264.128A2-17.615-12.1385.458A2-17.077-12.9054.338A2-15.560-13.0014.459A2-17.464-12.2573.017A3-12.247-13.6806.175A3-14.950-11.9174.932A3-13.517-11.8815.196A3-13.159-12.8596.315A3-13.085-10.4665.567A4-13.885-12.759了，425A4-13.661-13.6208.579A4-14.641-13.2859.702A4-13.771-15.0868.195A4-12.959-15.9058.628A5-14.778-15.4097.385A5-14.978—16.7766.915A5-16.217-16.8656.040A5-13.748-17.2656.160A5-13.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.70113.532_8.6441.0026.73cATOM31CBALAA4一14.62113.187_9.8021.0027.09cATOM32CALAA4-13.82215.011-8.2871.0028.76cATOM330ALAA4-13.05015,839_8.7691.0030.280ATOM34NALAA5—14.79715.334—7.4441.0027.61NATOM35CAALAA5-15.00516.708一7.0021.0032.20cATOM36CBALAA5-16.25916.808-6.1451.0028.22cATOM37CALAA5-13.79017.223-6.2381.0033.93cATOM380AUA5-13.45118.405-6.31410028.580ATOM39NVALA6-13.13716.330一5.50310033.08NATOM40CAVALA6-ll.93416.685_4.76410033.44cATOM41CBVALA6-11.53015.576-3.76910033.38cATOM42CGIVALA6-10.18515.894_3.13610035.97cATOM43CG2VALA6-12.59915.404-2.70210029.20cATOM44CVALA6-IO.78316.933_5.73010030.85cATOM450VALA6-10.05917.920—5.61010026.620ATOM46NSERA7—10.62616.032—6.69410031.55NATOM47CASERA—9.57716.161-7.69810031.70cATOM48CBSERA7-9.62514.980一8,66510035.87cATOM49OGSERA了—9.58913.750-7.96110040.860ATOM50CSERA-9.72317.467一8.46810027.45cATOM510SERA-8.73518.129-8.78010038.290ATOM52NSERA8-10,96317.831一8.77510031.44NATOM53CASERA8-11.24319.072_9.48710031.86cATOM54CBSERA8-12.73719.182—9.79610027.08cATOM55OGSERA8-13.02520.368-10.51610029.580ATOM56CSERA8-10.78720.290_868810030.02cATOM570SERA8-10.16021.201-9.23310031.630ATOM58NALAA9-11.10820.304_7.39710033.05NATOM59CAALAA9—10.72621.408一6.52310028.48cATOM60CBALAA9-11.37221.248_5.1481003484ATOM61CALAA99.20821.501-6.3941.0026.20ATOM620ALAA9-8.64022.593-6.4191.0029.65ATOM63NLYSA10—8.55620.350-6.2611.0025.05ATOM64CALYSA10-7.10020.306-6.2061.0028.57ATOM65CBLYSA10—6.603.18.872—6.0101.0031.94ATOM66CGLYSA10—6.84418.295-4.62010028.80ATOM67CDLYSA丄O-6.31716.867-4.5321.0028.88ATOM68CELYSA10-6.38716.320-3.1161.0030.92ATOM69NZLYSA10_5.87914.920-3.0461.0031.41ATOM70CLYSA10-6.51220.884-7.4881.0031.65ATOM710LYSA10—5.60821.717-7.4491.0028.85ATOM72NARGA11-7.04020.432-8.62110029.76ATOM73CAARGA11-6.58720.891-9.9261.0027.65ATOM74CBARGA11-7.34420.156—11.0331.0029.19ATOM75CGARGA11-7.14920.749-12.4171.00牧58ATOM76CDARGA11-7.95419.979-13.4551.0057.40ATOM77NEARGA11一9.37419.940—13.1071.0040.25ATOM78CZARGA11-10.24020.908-13.3941.0053.81ATOM79NH1ARGA11-9.83521.992-14.0411.0052.92ATOM80NH2ARGA11-11.51220.79613.0371.0049.16ATOM81CARGA11—6.76222.398—10.0771.0029.84ATOM820ARGA11—5.85023.095-10.5221.0030.08ATOM83NSERA12一7.93722.柳-9.7061.0033.81ATOM84CASERA12-8.21324.326-9.7961.0033.67ATOM85CBSERA12—9.66724.629_9.4231.0033.93ATOM86OGSEKA12-10.53424.397—10.5201.0041.64ATOM87CSERA12-7.25725.138_8.9261.0032.05ATOM880SERA12—6.78726.200-9.3321.0032.31ATOM89NLEUA13—6.96524.630-7.7331.0028.35ATOM90CALEUA13_6.07625.324-6.8081.0031.76ATOM91CBLEUA13-6.14224.690-5.4171.0028.56ATOM92CGLEUA13一5.45725.466-4.2891.0029.23ATOM93CD1LEUA13-5.85226.939-4.3131.0025.00ATOM94CD2LEUA13-5.77824.843-2.9411.0024.99ATOM95CLEUA13—4.63925.340-7.3181.0024.38ATOM960LEUA13-3.93026.332-7,1601.0028.27ATOM97NARGA14-4.21224.238-7.9281.0025.28ATOM98CAARGA14-2,86824.151-8.4881.0029.22ATOM99CBARGA14—2.60722.756-9.0631.0026.33ATOM100CGARGA14—2.54621.653_8.0151.0037.22ATOM101CDARGA14一l.98020.368_8,5991.0032.80ATOM102NEARGA14_2.03819.264-7.6441.0033.51ATOM103CZARGA14-2.94818.296_7.6721.0040.03ATOM104NH1ARGA14_3,88218.282_8.6141.0036.84ATOM105NH2ARGA14_2.92117.336_6.7591.0035.60ATOM106CARGA14_2.64425.218-9.56010031.22ATOM1070ARGA14-1.62325.904-9.56210028.32ATOM108NGLYA15-3.60825.354-10.46510034.73ATOM109CAGLYA15-3.53526.358-11.51010032.22ATOM110CGLYA15-3.47627.756-10.92910033.44ATOM1110GLYA15-2.68328.592-11.36310037.08ATOM112NGLUA16一4.32328.008_9.93710030.90ATOM113CAGLUA16—4.34029.292_9.24710040.01ATOM114CBGLUA16一5.49729.338-8.25010031.39ATOM115CGGUJA16—5.37330,419—7.19210044.27ATOM116CDGLUA16_6.53130.403一6.21310076.03ATOM1170E1GLUA16-7.57929.808—6.54410065.38ATOM1180E2GLUA16-6.39630.983_5.11510066.94ATOM119CGLUA16_3.02029.548_8.5291004349ATOM1200GLUA16-2.49930.664一8.54510037.47ATOM121NLEUA17-2.48228.505—7.90510037.86ATOM122CALEUA17-l.22828.611-7.1661003269ATOM123CBLEUA17一l.01627.3746.2881003101ATOM124CGLEUA17_1.90427.272—5.0461003679ATOM125CD1LEUA17_1.77025.902-4.4001003400ATOM126CD2LEUA17_1.57128.378-4.0541003336ATOM127CLEUA17—0.04128.800—8.1001003734CCONCCCCCNCONCCCCNCNNCONCCOCONCCCCCCONCCCCNCNNCONCCONCCCCOOCONCCCCCCATOM128ATOM129ATOM130ATOM131ATOM132ATOM133ATOM134ATOM135ATOM136ATOM137ATOM138ATOM139ATOM140ATOM141ATOM142ATOM143ATOM144ATOM45ATOM146ATOM147ATOM148A丁OM]49ATOM150ATOM151ATOM152ATOM153ATOM154ATOM155ATOM156ATOM157ATOM158ATOM159八TOM160「m5lATOM161LU"D」ATOM162ATOM163ATOM164ATOM165ATOM166ATOM167ATOM168ATOM169ATOM170ATOM171ATOM172ATOM173ATOM174ATOM175ATOM176ATOM177ATOM178ATOM179ATOM180ATOM181ATOM182ATOM183ATOM184ATOM185ATOM186ATOM187ATOM188ATOM189ATOM190ATOM191ATOM192ATOM193ATOM194ATOM195A170.85129.601-7.8241.0036.66A18—0.02828.065■-9.2081.0036.00A181.06728.174-10.1661.0037.03A180.99327.071-11.2231.0029.44A181.10425.670-10.6571.0043.79A181.93024.767-11.5571.0064.46A183.41725.048-11.3931.0072.85A184.25624.044-12.1051.0054.36A181.09429.544-10.8331.0038.69A182.16230.069—11.1431.0048.34A19—0.08330.118-11.0581.0037.19A19_0.16731.459-11.6191.0036.50A19-1.61831.864-11.8261.0038.43A190.54432.452-10.7051.0050.52A191.37233.244—11.1561.0046.79A200.22132.398-9.4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951.0048.541583CBGLUA1986.80635.0777.2891.0043.521584CGGLUA1985.36635.0957.7801.0049.661585CDGLUA1985.23835.6149.1941.0062.941586OE1GLUA1985.77836.7049.4791.0076.051587OE2GLUA1984.58934.93510.0181.0064.331588CGLUA1988.03535.5975.1491.0061.0615890GLUA1989.22035.2915.2761.0075.471590NGLYA1997.62636.6564.4551.0069.561591CAGLYA1998.55137.5283.7551.0063.981592CGLYA1999.55738.1914.6761.0050.0015930GLYA1999.68239.4174.6951.0076.191594GLYA1991595NINIC213.0862.397■5.1941.0034.901596MGMGC312.8347.149-13.9801.0068.271597NINIC4-9.56826.1198,3161.0061.901598MGMGC713.20829.982_6.6511.0055.891599MGMGC87.3775.445-2.0771.0051.811600MGMGC922.86817.547-15.20308448.781601MGMGC11-6.35730.545-0.3841.0052.781602MGc111603PP04B14.38615.122_4.67710043.52160401P04B15.80614.730_5.01210063.21160502P04B14.34416.590-4.33110059.90160603P04B13.51914.881_5.88710055.06160704P04B13.86414.312-3.51310034.421608P04B11609MGMGD114.3265.6101.14710022.161610MGMGD2-1.41615.580—0.68210053.081611PBADPD302-0.61915.432_4.7911004220161201BADPD302-l.08017,013—4.7971004359161302BADPD3020.89915.305_5.4181003588161403BADPD302-0.60814.928-3.42010041061615PAADPD302_2.78213.562—5.0221005374161601AADPD302-2.73513.709_3.3821006313161702AADPD302-4.29313.933-5.5641005783161803AADPD302-l.68714.580-5.7141005027161905'ADPD302_2.45412.183-5.38210059861620ADPD3021621MGMGE1-4.30215.9734.89810057891622MGE11623MGMGF114.72415.20215.4581005958ATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMTERATOMTERATOMMMMMMMMMMMMMMMMMMMMMMMMMM加加加MM加S6666SS6RSSSSOOSROOOOOROOOOOOOOOOOTTTTTTTTETTTTTTTETTTTTETTTTTTTTTTTAAAAAAAATAAAAAAACTAAAAATAAAAAAAAAAA91<table>tableseeoriginaldocumentpage92</column></row><table>證扁16920HOHS6618.37025.197-7.3101.0051.580HETATM16930HOHS6717.68510.292-15.5241.0058.270HETATM16940HOHS6812.64816.93219.9531.0063.670HETATM16950HOHS694.88038.339-5.1701.0060.940HETATM16960匪S7030.02112.8553.7351.0048.020HETATM16970HOHS7114.11139.633-11.1561.0064.770HETATM16980HOHS7227.0218.157-1.0671.0042.970HETATM16990■s7318.9358.79020.8631.0054.900HETATM17000HOHs7424.1856.910-1.2991.0045.770HETATM17010HOHs7515.3871.164-5.0261.0042.030HETATM17020HOHs768.47718.645-5.9371.0043.470HETATM17030HOHs778.3632.578-10.5591.0056.030HET扁17040腦s783.18136.918-6.8501.0056.060HETATM17050HOHs79-2.02116.9243.0491.0040.750HETATM17060HOHs8020.54122.038-7.6991.0050.640HETATM17070,s818.204-O.9372.4591.0042.920HETATM17080HOHs823.84021.262—11.7691.0051.240HETATM17090HOHs8313.24638.061-13.2291.0053.930HETATM17100■s8424.3935.304-13.5561.0056.210HETATM17110HOHs85—1.75428.341-14.3321.0047.120HETATM17120HOHs861.70214.012-2.5071.0040.630HETATM17130■s87-2.97516.490-3.0061.0040.640HETATM17140HOHs8816.8031.805-7.9881.0054.360HETATM17150HOHs8917.46721.749—20.1161.0054.420HETATM17160HOHs90-5.94027.5036.7331.0044.080HETATM17170■s917.23623.881-1.5141.0045.130HETATM17180■s92—0.13919.52511,7271.0049.040HETATM17190HOHs93_8.17929.545-2.3651.0064.320HETATM17200HOHs9415.69424.678-12.8991.0058.060HETATM17210HOHs9513.6479.99214.8691.0060.680HETATM17220腦s96-l.64710.621-2.4871.0052.230HETATM17230HOHs9715.7561.9917.9821.0052.310HETATM17240■s9823.32020.324-16.1091.0060.970HETATM17250HOHs993.753-2.5432.9921.0059.020IIETATM17260HOHs10025.9413.0543.5671.0049.410HETATM17270HOHs101-3.00310.116-6.8541.0053.540HETATM17280HOHs1026.379-3.0343.5101.0046.170HETATM17290HOHs1035.39713.78710.3261.0053.370HETATM17300HOHs104_8.67424.9546.1681.0049.770HETATM17310HOHs10518.4059.959-12.9881.0062.470HETATM17320HOHs10619.61522.180-16.5601.0050.980HETATM17330■s10712.77815.604-10.5901.0038.560HETATM17340HOHs1082.05314,7862.0511.0039.550HETATM17350s109—8.49516.67911.1071.0061.640HETATM17360HOHs110-4.64411.809-2.1971.0052.300HETATM17370鵬s11118.0603.814-9.2721.0048.120HETATM17380HOHs11224.25615.79514.9141.0048.680HETATM17390■s11322.07715.85316.8521.0058.750HETATM17400HOHs114-11.62327.3647.3501.0058.930HETATM17410HOHs11525.68929.034-6.2131.0062.100HETATM17420,s116-15.36620.610-7.2141.0057.380HETATM17430HOHs11727.44626.3829.7841.0052.130HETATM17440■s118一O.7806.946_1.2151.0067.580HETATM17450■s11932.40714.4163.9411.0061.020HETATM17460HOHs120_3.18014.315-IO.1761.0049.300HETATM17470HOHs121-13.63623.435—6.9821.0055.460HETATM17480HOHs12216.12326.725-9.2261.0062,320HETATM17490HOHs123—5.79113.788_7.7941.0054.450HETATM17500■s12416.36215.222_6.5161.0024.410HETATM17510■s12512.44615.9371.0561.,0025.060HETATM17520腳s12614.56517.219一5.2971..0024.180HETATM17530HOHs12716.9365.592-11.1621.0043.070HETATM17540HOHs12814.6765.669—9.8911.0051.760HETATM17550HOHs130-9.54212.446-10.6041.0064.680HETATM17560HOHs13112.96924.225-12.3561..0045.160HETATM17570HOHs13319.39729.024—0.5951.,0060.320HETATM17580■s13432.30111.柳2.6161,■0068.250HETATM17590HOHs1351.87715.851一12.7541..0056.83093表5為人源5，10-次甲基四氫葉酸合成酶與N5_亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體的坐標(biāo)。表5-8.49020.4807.2511.0048.11-2.08213.1198.0061.0052.530.00334.7674.9691.0044.34-12.84223.141一9.7161.0056.1610.21725.03516.8241.0059.342.8862.2254.5301.0062.0122.88828.011—7.3551.0064.535.94824.677-5.9451.0046.648.07830.204-10.6231.0071.164.00614.855-11.6731.0030.008.54115.171-9,9461.0030.001.43714.27011.3711.0030.00-4.14921.62113.5861.0030.00-9.97227.847-6.2001.0030.005.35932.01310.3721.0030.0031.2256.3801.4421.0030.000.83822.59612.1511.0030.00HETATM17600HETATM17610HETATM17620HETATM17630HETATM17640HETATM17650HETATM17660HETATM17670HETATM17680TER1769ATOM17700ATOM17710ATOM17720ATOM17730ATOM17740ATOM1775MGATOM1776NiATOM1777MGEND-9<u1_-344789<u1114445HHHHHHHHHHHHHHHiOOOOOOOOOOOOOOOGNGHHHHHHHHHHHHHHHMM941NMETA工16.3439.3557.94510034.02NATOM2CAMETA117.27710.5138.14310034.81CATOM3CBMETA11873310.0558.13710034.30CATOM4CGMETA119.3619.9369.53310034.35cATOM5SDMETA119.65111.53210.34610037.39sATOM6CEMETA120.36412.5219.02510032.27cATOM了CMETA117.08211.6297.12410035.19cATOM80META116.90012.7997.49410035.270ATOM9NALAA217.13411.2685.84510034.49NATOM10CAALAA216.72212.1714.77910034.40cATOM11CBALAA217.02811.5653.4291.0034.16cATOM12CALAA215.22612.4084.9211.0035.41cATOM130ALAA214.73213.5044.6341.0035.550ATOM14NALAA314.52311.3645.3761.0035.44NATOM15CAALAA313.08411.4055.6281.0035.26cATOM16CBALAA312.56110.0005.9641.0034.46cATOM17CALAA312.77912,3906.7591.0035.66cATOM180ALAA311.78113.1186.7201.0036.160ATOM19NALAA413.65912.4217.7551.0035.37NATOM20CAALAA413.51713.3328.8751.0035.04cATOM21CBALAA414.45412.93310.Oil1.0034.53cATOM22CALAA413.66614.8188.4871.0035.13cATOM230ALAA412.82915.6238.8741.0035.350ATOM24NALAA514.68615.1857.7061.0035.28NATOM25CAALAA514.85916.6017.2741.0035.00cATOM26CBALAA516.17416.7946.5561.0033.55cATOM27CALAA513.70817.1016.4011.0035.80cATOM280ALAA513.35118.2816.4471,0035.620ATOM29NVALA613.15716.1865.5971.0035.95NATOM30CAVALA611.90316.3904.8521.0036.31cATOM31CBVALA611,56715.1473.9391.0036.61cATOM32CGIVALA610.24515.3323.2251.0036.90cATOM33CG2VALA612.67514.8792.9121.0036.14cATOM34CVALA610.71416.7175.7991.0036.34cATOM350VALA69.90217.6025.5011.0036.520ATOM36NSERA710.62516.0146,9351.0036.23NATOM37CASERA了9.54316.2337.9161.0036.04cATOM38CBSERA79.36515.0188.83110035.80cATOM39OGSERA79.02213.8698.0761.0035.560ATOM40CSERA79.75717.4948.7431.0035.72cATOM410SERA78.79418.1529.1471.0035.800ATOM42NSERA811.02217.8169.0011.0035.39NATOM43CASERA811.38119.1039.5701.0035.29cATOM44CBSERA812.90419.2459.6451.0035.24cATOM45OGSERA813.28220.34110.46010034.910ATOM46CSERA810.76720.1828.6811.0035.40cATOM470SERA89.95620.9819.1441.0035.460ATOM48NALAA911.12420.1387.3931.0035.75NATOM49CAALAA910.65921.0736.3561.0035.63cATOM50CBALAA911.17720.6504.99910035.19cATOM51CALAA99.15121.2076.31310035.74cATOM520ALAA98.62222，3236.23610035.860ATOM53NLYSA108.46420.0666.3591.0035.72NATOM54CALYSA107.00720.0426.37310035.34cATOM55CBLYSA106.47518.6076.18910035.89cATOM56CGLYSA106.743.17.9754.80310035.39cATOM57CDLYSA105.95916.6874.61210035.34cATOM58CELYSA106.74115.6813.78210036.74cATOM59NZLYSA105.95115.1242.61610038.06NATOM60ATOM61ATOM62ATOM63ATOM64ATOM65ATOM66ATOM67ATOM68ATOM69ATOM70ATOM71ATOM72ATOM73ATOM74ATOM75ATOM76ATOM77ATOM78ATOM79ATOM80ATOM81ATOM82ATOM83ATOM84ATOM85ATOM86ATOM87ATOM88ATOM89ATOM90ATOM91ATOM92「ni^lATOM93LUIOJ」ATOM94ATOM95ATOM96ATOM97ATOM98ATOM99ATOM100ATOM101ATOM102ATOM103ATOM104ATOM105ATOM106ATOM107ATOM108ATOM109ATOM110ATOM111ATOM112ATOM113ATOM114ATOM115ATOM116ATOM117ATOM118ATOM119ATOM120ATOM121ATOM122ATOM123ATOM124ATOM125ATOM126ATOM12735.06C35.05035.28N35.05C35.16C35,03C35.48C35.57N35.25C34.56N34.61N34.75C34.50034.69N35.12C35.50C35.99035.22C35.48035.06N34.71C34.75C34.22C33.98C34.48C34.80C34.83034.93N34.67C34.56C34,50C35.05C34.75N34.67C34.62N34.67N34.94C34，93035.07N35.17C35.50C35.64035.47N35,85C35.98C36.51C37.57C36,95038.34035.72C36.08035.69N34.96C34,57C34.35C33.87C34.13C35.08C35.49035,29N35,20C34.52C34.62C34.79C35.36C36.16N35.37C35.620106，46720.6927.659105,59221.5647.600117.01320.2918.808116,57720.80010,110117.36020.08911,220116-84420.27312.647117.98820.01013.620119.26820.38813.0001110.05421.39313,388119,73322.13714,44911.18021.63212.715116.70022.33610.197115.70523.03010.434127.91222.8529.985128.15924.2999.884129.49924.5889.1891210.58123.8949.794127.06224.9879.089126.28125.7709,635137,01224.6647.797136.07425.2576.843136.30524.6645.441135.71925.2904，165136.14026.7273.930136.14324,4522.996134.60425.1687.281133.82226.0596-969144,23724.1188.020142.90924.0468.652142.68322.6969,317142，謹(jǐn)21.5478,387141-81420.4578.716141,83219.4087.695142.53218.2807.785143.26918.0288.867142，48417,4006.794142.72125.1359,701141.65325.7249.800153.75425.38310.502153.67126.38111.553153.49427.72210,889152,48128,40711,080164.46928.08410'069164，40729.3619.378165,52329.4848.344165.62530.8757.731166.07630,8396.274166.90029,9565,896165.59931,7065.505163.04729.5628.717162.45730,6278.853172,56428.5268,021171.28128.5707.310171.08027.3636,386171.69927.3564.976171.66725.9494.389171.01528.3314,031170.10928.6978.25317一O.82329.4377.956180.14927.9949,38718_0,91428.12010,38818_0.84427.01011.42318—0.86725.62410，82118—1.21324.54111,S3618_2.71824.30111.86218—3.07922,84411.92518-o.87629.49911,05518一l.91929.99411.504SSGGGGGGGGGGGRRRRRRUUUUUUUUGGGGGGGGGGGYYYYUUUUUUUUUUUUUUUUUSSSSSSSSS1212121212ABGDEZHHABGABGDDABGDEZHHA:ACDGDEEABGDDABGDENICONCCCCNCNNCONCCOCONCCCCCCONCCCCNCNNCONCCONCCCCOOCCDNCCCCCCONCCCCCNCOoooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo<table>tableseeoriginaldocumentpage97</column></row><table>ATOM196CZARGA29-13.94729.3316.5771.0031.72CATOM197NH1ARGA29-15.00229.9227.1251.0031.81NATOM198NH2ARGA29-13.79528.0176.6911.0031.46NATOM199CARGA29-12.99234.1663.8451.0033.79CATOM2000ARGA29-12.93633.3592.8901.0033.520ATOM201NLEUA30-14.10034.8614.1461.0033.80NATOM202CALEUA30-15.33434.7533.3431.0033.45CATOM203CBLEUA30-16.52835.4144.0331.0033.67CATOM204CGLEUA30-16.99534.7855.3491.0034.30cATOM205CD1固A30-17.73235.8106.2181.0034.57cATOM206CD2LEUA30-17.84833.5315.1381.0033.97cATOM207CLEUA30-15.17535.3281.9431.0033.20cATOM2080LEUA30-15.65334.7270.9781.0033.370ATOM209NARGA31-14.50436.4781.8371.0033.10NATOM210CAARGA31-14.30437.1610.5521.0032.72cATOM211CBARGA3113.49038.4550.7361.0032.81cATOM212CGARGA31-13.07539.106_0.5821.0032.51cATOM213CDARGA31-12.08840.231_0.4041.0032.09cATOM214NEARGA31-IO.70639.773-O.2301.0031.96NATOM215CZARGA31-9.89639.375—1.2111.0031.81cATOM216NH1ARGA31-10.32139.340-2.4731.0030.32NATOM217NH2ARGA31-8.65338.985-0.9111.0031.88NATOM218CARGA31-13.63036.243-0.4831.0032.20CATOM2190ARGA31—14.15736.012-l.5801.0032.240ATOM220NGLNA32-12.46835.719_0.1021.0031.98NATOM221CAGLNA32-ll.63734.893-o.9641.0031.72CATOM222CBGLNA32-10.33334.572-o.2331.0031.66cATOM223CGGLNA32-9.05735.028-o.9251.0031.17cATOM224CDGLNA32-7.85334.958-l.0001.0031.20cATOM2250E1GLNA32—7.80235.6431.0171.0031.960ATOM226NE2GLNA32一6.87734.137-o.3531.0030.78NATOM227CGLNA32-12.38833.611-l.3381.0031.61cATOM2280GLNA32-12.18533.072-2.4251.0031.040ATOM229NSERA33-13.25833.150-O.4321.0031.90NATOM230CASERA33-14,14132.004-O.6601.0031.20cATOM231CBSERA33-14.78631.5450.6451.0031.24cATOM232OGSERA33-13.81231.0931.5641.0031.100ATOM233CSERA33-15.21732.325_1.6871.0031.35cATOM2340SERA33-15.45131.541-2.6061.0031.350ATOM235NARGA34-15.86933.477-1.5511.0031.68NATOM236CAARGA34-16.77433.965-2.6151.0031.88cATOM237CBARGA34-17.30435.361-2.2871.0033.05cATOM238CGARGA34-18.80735.447-2.0741.0034.15cATOM239CDARGA34-19.19336.861-l.6211.0035.19cATOM240NEARGA34-19.13936.989-0.1561.0034.99NATOM241CZARGA34-19.11538.1410.5141.0035.64cATOM242NH1ARGA34-19.12439.314_0.1361.0036.01NATOM243NH2ARGA34-19.06638.1171.847〗.DO35.58NATOM244CARGA34-16.12933.990-4.0141.0031.50CATOM2450ARGA34-16.81833.780_5.0141.0031.730ATOM246NVALA35-14.81634.230_4.0831.0031.20NATOM247CAVALA35-14.11634.312_5.3761.0030.76CATOM248CBVALA35-12.88135.246_5.3141.0030.82cATOM249CGIVALA35-12.25635.404—6.6761.0030.70cATOM250CG2VALA35-13.29636.608-4.8171.0031.39cATOM251CVALA35-13.78932.936—5.9941.0030.92cATOM2520VALA35-14.09732.690-7.1721.0031.250ATOM253NLEUA36-13.19532.038_5.2011.0030.91NATOM254CALEUA36-12.95030.651-5.6321.0030.38cATOM255CBLEUA36-12.19029.870一4.5631.0029.61cATOM256CGLEUA36-10.66529.920—4.6041.,0029.83cATOM257CD1LEUA36-10.15231.188_3.9851,,0030.58cATOM258CD2固A36-10.12428.764_3.8381.,0029.69cATOM259CLEUA36-14.23729.918-5.9841.,0030.04cATOM2600LEUA36-14.28329.173—6.9591.,0030.200ATOM261NSERA37-15.27830.132_5.1881.0029.70INATOM262CASERA37-16.58629.573一5.4851..0030.05cATOM263CBSERA37—17.64130.243—4.6271,,0030.58c<table>tableseeoriginaldocumentpage99</column></row><table>ATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOM-16.65922.497-11.3531.0027.5917.90922.461-10.5041.0027.27-18.02021.579-9.4261.0026.98-19.17021.529-8.6501.0026.70-20.23322.341-8.9741.0027.25-21.38922.277-8.2331.0027.56-20.15523.215-10.0441.0027.41-18.99723.270-10.8011.0027.46-17.76621.030-13.1001.0027.98-18.60020.159-12.8351.0028.26-17.87721.807-14.1701.0028.09-19.06921.731-15.0121.0028.04-18.98522.749—16.1481.0027.93-19.43524.116-15.7151.0028.12-19.00425.207-16.6751.0029.62-18.47324.946-17.7781.0029.29-19.22526.461-16.2611.0029.92-19.34520.319-15.5331.0027.79-20.44019.799-15.3511.0027.66-18.33319.704-16.1441.0028.11-18.42418.357-16.7381.0027.88-17.16818.063-17.5801.0027.86-16.98019.055-18.7561.0028.32-15.67718.814-19.5291.0028.63-15.16520.102-20.2091.0029.11-15.33320.238-21.7051.0028.63-18.69017.207-15.7571.0027.78—19.24716.192-16.140.0027.80-18.32817.372-14.4891.0028.24-18.30416.240-13.5531.0027.62-17.17216.395-12.5311.0027.65-17.63317.077一ll.3721.0028.14-19.64015.932-12.8621.0027.41-20.47816.807-12.6661.0027.27-19.80114.668-12.4771.0027.67-21.06914.142-12.0021.0027.32-21.54113.041-12.9531.0027.62-21.68813.570-14.3931.0028.12-21.53812.509—15.4951.0028.41-21.71913.158-16.8821.0028.76-20.510IS-929-17.3071.0028.24-20.942IS.642一IO.5731.0027.51-21.69914,074-9.6841.0027.91-19.98512.744一IO.3281.0027.35-19.77212.245-8.9601.0027.07-19.58610.726_8.9401.0026.87-20.5919.980-9.8391.0026.65-20.2018.523-10.1161.0026.31-19.7797.843_8.9021.0026.34-20.6077.346-7.9911.0026.62-21.9257.414-8.1551.0026.60-20.111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871002989940CAVALA119一243122.5566.5051002990941CBVALA119-250801.2126.2481003036942CGIVALA119-24.6480.5884.9201002983943CG2亂A119—248690.2127.3951003023ATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMA，ATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATOMATATATATATATATATATATATOMATOMCCCONCONCCCCCCONCCCCCONCCCCONCONCCCCCONCCONCCCCOOCONCCONCCCOOCONCCCCoooooooooo108<table>tableseeoriginaldocumentpage109</column></row><table>ATOM1012CGLEUA129-15.29110.133-6.0181.0026.49CATOM1013CD1LEUA129-14.34610.143-4.8331.0026.53Catom1014cd2leua129-16.47311.083-5.7961.0026.69cATOM1015CLEUA129-14.63510.508-9.7881.0027.38CATOM10160LEUA129-13.52510.200-10.2781.0027.680ATOM1017NASPA130一15.45611.412-IO.3261.0027.22Natom1018caaspa130—15.06412.268—11.4351.0027.33catom1019cbaspa130-16.30712.844-12.1321.0027.18catom1020cgaspa130-17.04911.788-12.9681.0027.27catom1021od1aspa130-16.38011.090-13.7601.0027.460ATOM1022OD2ASPA130-18.29311.648-12.8271.0027.150atom1023caspa130-14.09213.366-io.9591.0027.51cATOM10240ASPA130-13.02113.579-11.5771.0027.530ATOM1025NLEUA13143814.009-9.834.0027.14NATOM1026CALEUA131-13.65015.123-9.2831.0026.88cATOM1027cbLEUA131—14.15616.449-9.8601.0026.70cATOM1028CGLEUA131—13.73717.705-9.0961.0026.61cATOM1029CD1LEUA131-12.45518.277-9.6431.0027.04cATOM1030CD2LEUA131-14.82318.726-9.1941.0026.99catom1031cleua131-13.583is-193-7.7321.0026.61catom10320leuA131一14.606is.186-7.0291.0026.340ATOM1033nILEA132-12.37115.295-7.2071.0026.84NATOM1034CAILEA132-12.17415.461-5.7801.0026.38cATOM1035CBILEA132一ll.13014.454-5.3111.0026.55catom1036CGIilea132—11.53213.052-5.7891.0026.01cATOM1037CD1ILEA132-10.43212.039-5.8061.0026.14cATOM1038CG2ILEA132-10.92414.535-3.7781.0026.77cATOM1039CILEA132-11.73116.907-5.4901.0026.41cATOM10400ILEA132-10.79017.410-6.1141.0026.830atom1041nphea133-12.42517.610-4.5981.0026.15nATOM1042CAPHEA13392118.922-4.1911.0026.40cATOM1043CBPHEA133-13.0。719.860-3.7241.0026.17cATOM1044CGPHEA133-13.78320.468-4.8251.0026.08cATOM1045CD1PHEA133-13.33621.607-5.4491.0026.22cATOM1046CE1PHEA13314.05722.179-6.4631.0026.32cATOM1047CZPHEA133-15.25421.625-6.8621.0026.06cATOM1048CE2PHEA133-15.72520.496-6.2481.0025.94cATOM1049CD2PHEA133-14.98519.916-5.2261.0026.24cATOM1050CPHEA13300618.641-3.0491.0026.96cATOM10510PHEA133-11.36417.937-2.0801.0026.990atom1052nmeta134一9.81119.185-3.1541.0027.26nATOM1053CAMETA134一8.77118.802-2.2281.0027.43catom1054cbmeta134-7.53418.437-3.0181.0027.86cATOM1055CGMET134-7.73917.292-3.983.0027.30cATOM1056SDMETA134-7.17715.734-3.2771.0028.01sATOM1057CEMETA134一5.53316.209-2.7231.0028.03cATOM1058CMETA134一8.46619.901-l.2101.0027.97catom10590meta134-8.27021.066-l.5891.0027.940atom1060nproa135一8.44319.5370.0891.0028.19natom1061CAPROA135一8.01520.4441.1401.0028.68cATOM1062CBPROA135一8.62519.8262.3901.0028.21cATOM1063cgPROA135-8.61018.4062.1261.0028.04cATOM1064CDPROA135一8.80518.2240.6441.0028.05cATOM1065CPROA135--6.49520.4011.2231.0029.60cATOM10660PROA135一5.85519.6730.4421.0029.450ATOM1067NGLYA136一5.93221.1742.1501..0029.83nATOM1068CAGLYA136一4.50621.1522.4081.0030.22catom1069Cglya136一4.08622.4293.0811.0031.27cATOM10700GLYA136一4.70523.4932.8971.0031.360ATOM1071NLEUA137—3.02622.3183.8681.0031.83NATOM1072CALEUA137-2.42523.4624.5151.,0031.87cATOM1073CBLEUA137_1.42923.0175.5911.■0032.17catom1074cgleua137—1.99322.5866.9321.,0032.14cATOM1075cd1LEUA137_0.85622.4067.8901.,0033.30cATOM1076CD2LEUA137-2.99923.6057.4911.0032.85cATOM1077cLEUA137_1.73724.3703.5031.,0031.92catom10780leua137—1.84225.5873.6201.■0032.210atom1079nglya138—1.02723.7972.5291,.0031.88n110ATOM1080CAGLYA138一O.39524.6171.4901.0032.02ATOM1081CGLYA138-O.19623.9410.1521.0032.07ATOM10820GLYA138-0.19322.7320.0471.0032.10ATOM1083NPHEA139-O.03724.734-0.8911.0032.65ATOM1084CAPHEA1390.25424.199-2.2141.0032.23ATOM1085CBPHEA139—1.00824.124-3.0601.0031.67ATOM1086CGPHEA139-2.17523.497-2.3661.0031.61ATOM1,CD1PHEA139-2.47722.157-2.5671.0031.31ATOM1088CE1PHEA139-3.56421.565-l.9311.0030.63ATOM1089CZPHEA139一4.35422.309-l.0791.0030.51ATOM1090CE2PHEA139一4.06923.6590.8711.0030.79ATOM1091CD2PHEA139-2.98724.249-l.5151.0031.24ATOM1092CPHEA1391.19125.176-2.8761.0033.04ATOM10930PHEA1391.11226.3762.6081.0033.56ATOM1094NASPA1402.07824.682-3.7361.0033.22ATOM1095CAASPA1402.83825.567-4.6091.0033.38ATOM1096CBASPA1404.33925.2184.5731.0033.74ATOM1097CGASPA1404.61623.767-4.9381.0033.81ATOM1098OD1ASPA403.73423.140-5.5751.0033.68ATOM1099OD2ASPAMO5.70323.244一4.5871.0033.92ATOM1100CASPA1402.25025.529■-6.0351.0033.27ATOM11010ASPA1401.43024.669-6.3621.0032.94ATOM1102NLYSA1412.66126.470-6.8791.0033.62ATOM1103CALYSA1412.20626.507-8.2681.0033.37ATOM1104CBLYSA1412.60227.852-8.8981.0033.42ATOM1105CGLYSA1411.94229.074-8.2261.0033.50ATOM1106CDLYSA1412.41830.417-8.7981.0033.27ATOM1107CELYSA1411.80631.583-8.0291.0033.54ATOM1108NZLYSA1412.35932.908-8.4381.0033.78ATOM1109CLYSA1412.70025.302-9.1111.0033.15ATOM11100LYSA1412.10424.972-10.1271.0032.02ATOM1111NHISA1423.80324.675-8.6741.0034.15ATOM1112CAHSA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m16540atom16550atom16560atom16570atom16580atom16590atom16600atom16610atom16620atom16630atom16640atom16650atom16660atom16670atom16680atom16690atom16700atom16710atom16720atom16730atom16740atom16750atom16760atom16770atom16780atom16790atom16800atom16810atom16820atom16830atom16840atom16850atom16860atom16870atom16880atom16890atom16900atom16910-21.7738.266-o.65518.33222.365-8.123-22.8685.869-l.0328.73527.52014.2945.56132.40910.7303.48520.878—11.25614.73415.15315.7052.35438.9845.1333.60414.2608.6906.11724.5585.825-5.24228.0896.696—0.0228.022-3.40116.柳13.96616.6987.58023.905-l.5970.19819.80111.67517.8789.768—14.8016.42613.83013.793一6.8139.8852.462-15.09920.665-6.48212.63815.9641.07816.42015.111-6.5877.0267.446-3.404-7.74327.7928.4630.84915.057-5.06915.2791.219-5.1146.0193.368-3.7585.65527.51811.523-20.69810.004_2.59713.9333.010-6.968—8.81725.3286.5087.396-0.2404.7821.08620.398-io.73116.08744,3110.072-l.63815.412-o.177-2.38816.724-3.45218.19127.7661.43512.14731.009—6.80019.11710.785-13.09326.10420.028-14.05427.61413.07912.159_9.65727.743_5,754-16.78220.916_8.8066.35228.700一16.7758.74122.83271924.76228.026-7.3998.88323.287is-95510.93816.974is.40622.0730.3551.830-10.07212.611-4.68018.24925.647-7.442—0.71315.155-2.57819.70831.32011.66815.94926.86020.6118.70425.78816.3372.22727.63610.490_9.74726.82110.81613.61740.4493.6405.8864.60114.75821,1880.167-9.35710.89911.079-17.56819.0392.484一9.7578.06430.374-11.2702.43824.92610.90523.9206.1577.13411.31625.13217.1040.71023.33512.49220.55822.652一5.20316.9951.869—8,0321.0043.4401.0046.4201.0042,9001.0054，4401.0049.1701.0024.6801.0049.2101.0063.3601.0044.7501.0036.2901.0028.8101.0048.2301.0044.5901.0035.1901.0042.0201.0046，8901.0040.6201.0048.450i.0036,0401.0014.8201.0015.3901.0018.9001.0030.8701.0024,8301.0026,1901.0050.0501.0040.0001.0034，2701.0024.8501.0035.9701.0045,1701.0030.3601.0046.5701.0042.0001.0036.9101.0033.2301.0049.7801.0036.8901.0038.1901.0042.920〗.0051.4601.00肌6601.0042.5401.0050.5901.0064.2201.00肌9701.0046.4901.0056,2001.0036，8001.0047.6901.0048.8201,0042.89。1.0053.4701.0052.0501.0048.1001.0045.3301.0043.0901.0046.1401.0058.9401.0053.9001.0052，1201,0051.1201.0029.4601.0036.1201.0043.3901,0047,9701,0051.8601.0042.320-2345678901234567890123456789012345678901--777777778888888888999999999900000000001H1.5舊附附HHH555舊附HHHHHHH!HHHHHHHHHHHHHHHHHHHHHHHHH附M附W附S附附SS州S州SS州州州S州刑州州州OO000OO000O00O0000000OO000OO00O00O00O0O0OO0O0OO000O0O00O0000O02O0202HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH143各種MTHFS突變體以及人野生型MTHFS的酶活性測(cè)定通過(guò)對(duì)各種MTHFS同源蛋白的生物化學(xué)和酶學(xué)性質(zhì)的研究，發(fā)明人挑選了15個(gè)位點(diǎn)進(jìn)行定點(diǎn)突變研究。突變使用的原理是通過(guò)使用定點(diǎn)突變的方法將分析得到的潛在關(guān)鍵位點(diǎn)進(jìn)行突變，突變的方法使用STRATAGENE公司定點(diǎn)突變?cè)噭┖?，通過(guò)PCR—步得到突變體，經(jīng)過(guò)DNA測(cè)序來(lái)驗(yàn)證突變是否成功。潛在的關(guān)鍵氨基酸殘基都被突變成為丙氨酸(A)，由于丙氨酸具有最簡(jiǎn)單的氨基酸支鏈，同時(shí)又對(duì)蛋白質(zhì)的二級(jí)結(jié)構(gòu)產(chǎn)生很小的影響，所以這一方法已經(jīng)成為廣泛用于定點(diǎn)突變中。以5-甲基四氫葉酸為底物，加入任一種MTHFS突變體或人野生型MTHFS以及輔助因子鎂離子、ATP(三磷酸腺苷)。室溫反應(yīng)2分鐘，通過(guò)紫外吸收光譜檢測(cè)360nm處吸收光強(qiáng)度的變化，確定生成物5，10-亞甲基四氫葉酸的含量，進(jìn)而確定酶活性的強(qiáng)弱。將人野生型MTHFS的活性設(shè)為100%，突變體酶的相對(duì)活性為與人野生型酶活性比較獲得。表7.MTHFS突變體的相對(duì)活性<table>tableseeoriginaldocumentpage144</column></row><table>人源5，10-次甲基四氫葉酸合成酶及其復(fù)合物的空間構(gòu)型<table>tableseeoriginaldocumentpage144</column></row><table>經(jīng)結(jié)構(gòu)修正得到人源MTHFS的原子坐標(biāo)，MTHFS的三維結(jié)構(gòu)見(jiàn)圖3，其中該酶的三維結(jié)構(gòu)中包括a螺旋和P片層，其位點(diǎn)如下a12-18;a226-42;a344-48;P150-54;a465-73;P277-84;P389-95;P4131-134;P5138-140;P6145-146;a5152-162;P7169-173;P8194-196。從突變實(shí)驗(yàn)以及三維結(jié)構(gòu)圖的結(jié)果可知以下幾個(gè)位點(diǎn)對(duì)于酶發(fā)揮活性是關(guān)鍵的F55、M58、E61、M90、Y152、Y153通過(guò)與底物5-甲基四氫葉酸中的蝶呤基團(tuán)結(jié)合來(lái)發(fā)揮酶活性；F85、W109、R148通過(guò)與底物5-甲基四氫葉酸中的對(duì)氨基苯甲酸基團(tuán)結(jié)合來(lái)發(fā)揮酶活性；KIO、R14、R145、D154通過(guò)與ATP結(jié)合來(lái)發(fā)揮酶活性。人源5，10-次甲基四氫葉酸合成酶與ADP復(fù)合物蛋白結(jié)合磷酸基團(tuán)以后，在結(jié)構(gòu)中原來(lái)無(wú)法建模的部分，E187-D189，在人源MTHFS結(jié)合ADP的復(fù)合物結(jié)構(gòu)中得到了很好的構(gòu)建。E187-D189位于ATP的結(jié)合部位，D189與ATP水解下來(lái)的磷酸的01氧原子形成一個(gè)氫鍵，并且結(jié)合了一個(gè)鎂離子。另外，D189和K10形成一個(gè)鹽橋。這些新的化學(xué)作用鍵的形成，使得在脫輔基蛋白結(jié)構(gòu)中無(wú)法建模的部分(A174-D193)得到了穩(wěn)定。由于在三維結(jié)構(gòu)中只能顯示ADP的兩個(gè)磷酸基團(tuán)和一個(gè)從ATP上水解下來(lái)的磷酸基團(tuán)，其中水解下來(lái)的磷酸將作為親核基團(tuán)參與酶反應(yīng)。此磷酸基團(tuán)位于a5(152-161位氨基酸)的一段，151-153位氨基酸殘基與之有相互作用。使用ATP類(lèi)似物，例如使用硫類(lèi)似物來(lái)替代磷酸基團(tuán)設(shè)計(jì)小分子藥物可以抑制此處磷酸基團(tuán)進(jìn)入酶分子，從而抑制酶活性。人源5，10-次甲基四氫葉酸合成酶與谷氨酸復(fù)合物結(jié)合在一個(gè)111-116位氨基酸殘基附近的中性電荷裂縫上，該裂縫為P3(89-95氨基酸區(qū)段)和134(131-135氨基酸區(qū)段)之間的一個(gè)大Loop環(huán)區(qū)域，其中G115和E116與它有4埃之內(nèi)的相互作用，谷氨酸部分通過(guò)一個(gè)圓形口突出結(jié)合口袋以外而位于蛋白表面。這種特異性的結(jié)合可以被用來(lái)設(shè)計(jì)多肽抑制劑，用來(lái)設(shè)計(jì)特異性的酶抑制劑。人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物氨基酸殘基F55，L56，S57，M58，E61，P81，Y83，M90，W109，P135，R148，K150，Y152和Y153圍成的一個(gè)大球形口袋中，5-亞胺磷酸反應(yīng)中間態(tài)的蝶呤環(huán)就位于球形的中間。4-螺旋和3、4-折疊，以及它們之間的長(zhǎng)loop，形成了一個(gè)帽子形狀的結(jié)構(gòu)覆蓋在活性中心的上面。W109位于蝶呤環(huán)的上面，并且看上去是用來(lái)關(guān)閉活性中心的蓋子，W109與蝶呤環(huán)的距離為7A,說(shuō)明兩者之間沒(méi)有相互作用。蝶呤環(huán)的構(gòu)象并不是一個(gè)平面，它在C6位置沿垂直軸方向伸展。61位的羧基氧與蝶呤環(huán)的N3和氨基氮相互作用。E61在直系同源MTHFS中是保守殘基。M90與蝶呤環(huán)的N8形成一個(gè)氫鍵。在蝶呤環(huán)和F55，Y83，P135，Y152，Y153的芳香側(cè)鏈之間存在著很多疏水的范德華力相互作用。Y83在野生型和復(fù)合物的晶體中區(qū)別最大，Y83的側(cè)鏈發(fā)生旋轉(zhuǎn)形成一個(gè)垂直狀態(tài)，使得蝶呤環(huán)進(jìn)入結(jié)合口袋。Y83側(cè)鏈的移動(dòng)，反過(guò)來(lái)使Y85的側(cè)鏈遠(yuǎn)離了結(jié)合口袋。所述的氨基酸殘基均為參與酶反應(yīng)的關(guān)鍵基團(tuán)，可以為設(shè)計(jì)小分子藥物提供結(jié)構(gòu)信息。此N5-亞胺磷酸過(guò)渡態(tài)中，蝶呤環(huán)以C4a與C8a之間的鍵為軸發(fā)生了扭轉(zhuǎn)，所述喋呤環(huán)上的電子發(fā)生了重排。人源5，10-次甲基四氫葉酸合成酶與生成物10-甲酰四氫葉酸的復(fù)合物與人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物空間的構(gòu)型總體上相同，只有Y83側(cè)鏈的移動(dòng)位置不同。Y83可以用來(lái)設(shè)計(jì)抑制其移動(dòng)的小分子，進(jìn)而設(shè)計(jì)小分子藥物。上面的結(jié)構(gòu)信息可用于設(shè)計(jì)或制造藥物。本發(fā)明還提供了5，10-次甲基四氫葉酸合成酶或其復(fù)合物的抑制劑，包括與上述的結(jié)構(gòu)域類(lèi)似或者與上述的結(jié)構(gòu)域結(jié)合的小分子化合物、短肽、多肽、核酸、抗體或免疫結(jié)合物。本發(fā)明還提供了用于移植癌癥的藥物組合物，包括5，10-次甲基四氫葉酸合成酶或其復(fù)合物的抑制劑，以及藥用載體或賦形物。序列表〈110〉中國(guó)科學(xué)院生物物理研究所〈120〉人源5，10-次甲基四氫葉酸合成酶及其復(fù)合物的結(jié)晶方法、晶體以及應(yīng)用〈130〉P25377IBP〈160>1〈170>PatentInversion3.2〈210>1〈211>203〈212>PRT〈213〉人類(lèi)〈400>1MetAlaAlaAlaAlaValSerSerAlaLysArgSerLeuArgGlyGlu151015LeuLysGinArgLeuArgAlaMetSerAlaGluGluArgLeuArgGin202530SerArgValLeuSerGinLysVallieAlaHisSerGluTyrGinLys354045SerLysArglieSerliePheLeuSerMetGinAspGlulieGluThr505560GluGlulielieLysAspliePheGinArgGlyLyslieCysPhelie65707580ProArgTyrArgPheGinSerAsnHisMetAspMetValArglieGlu859095SerProGluGlulieSerLeuLeuProLysThrSerTrpAsnliePro100105110GinProGlyGluGlyAspValArgGluGluAlaLeuSerThrGlyGly115120125LeuAspLeuliePheMetProGlyLeuGlyPheAspLysHisGlyAsn130135140ArgLeuGlyArgGlyLysGlyTyrTyrAspAlaTyrLeuLysArgCys145150155160LeuGinHisGinGluValLysProTyrThrLeuAlaLeuAlaPheLys165170175GluGinlieCysLeuGinValProValAsnGluAsnAspMetLysVal180185190AspGluValLeuTyrGluAspSerSerThrAla195200權(quán)利要求一種人源5，10-次甲基四氫葉酸合成酶的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a＝48.70±2b＝145.12±2c＝59.21±2α＝90.00°，β＝90.00°，γ＝90.00°。F2009101344744C0000011.tif,F2009101344744C0000012.tif,F2009101344744C0000013.tif2.—種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a=48.51±2A，b=145.22±2A，c=59.65±2A;a=90.00°，P=90.00°，y=90.00°。3.—種人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體，其特征在于所述晶體的空間群為P21212，晶胞參數(shù)為a=59.579±2A,b=144.101±2A，c=48.532±2A;a=90.00°，P=90.00°，y=90.00°。4.一種人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a=48.45±2A,b=145.23±2A，c=59.60±2A;a=90.00°，=90.00°，y=90.00°。5.—種人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a=48.69±2A,b=141.03±2A,c=59.99±2人；a=90.00°，P=90.00°，y=90.00°。6.—種人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體，其特征在于所述晶體的空間群為C2221，晶胞參數(shù)為a=48.86±2A,b=144.97±2A,c=59.41±2人；a=90.00°，P=90.00°，y=90.00°。7.根據(jù)權(quán)利要求1所述的人源5，10-次甲基四氫葉酸合成酶的晶體的結(jié)晶方法，包括將所述人源5，10-次甲基四氫葉酸合成酶的溶液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于，在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH二6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。8.根據(jù)權(quán)利要求2或3所述的人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP溶液，再加入還原劑和鎂離子，混合后得到人源5，10-次甲基四氫葉酸合成酶與ATP的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與ATP的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于，在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH二6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。9.根據(jù)權(quán)利要求4所述的人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入谷氨酸溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與谷氨酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與谷氨酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于，在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH二6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。10.根據(jù)權(quán)利要求5所述的人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP、5-甲酰四氫葉酸、巰基乙醇、鎂離子溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在1-2天后收集晶體；其特征在于，在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH二6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。11.根據(jù)權(quán)利要求6所述的人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體的結(jié)晶方法，包括向所述人源5，10-次甲基四氫葉酸合成酶的溶液中加入ATP、5-甲酰四氫葉酸、巰基乙醇、鎂離子溶液，混合后得到人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液；將所述人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的混合液與池液混合；利用懸滴法或坐滴法進(jìn)行結(jié)晶；在4-14天后收集晶體；其特征在于，在5至25攝氏度、50%至80%的濕度范圍內(nèi)，池液為含0.01-0.03mol/LMgCl2、0.01-0.04mol/LNiCl2、20%-30%PEG3000至4000、0.lmol/L的pH二6.4-7.0的N-2-羥乙基哌嗪-N-2-乙磺酸或嗎啉乙磺酸緩沖液。12.根據(jù)權(quán)利要求1所述的人源5，10-次甲基四氫葉酸合成酶的晶體的F55、M58、E61、190、￥152、￥153位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與底物5-甲酰四氫葉酸中的蝶呤基團(tuán)結(jié)合。13.根據(jù)權(quán)利要求1所述的人源5，10-次甲基四氫葉酸合成酶的晶體的F85、W109、尺148位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與底物5-甲酰四氫葉酸中的對(duì)氨基苯甲酸基團(tuán)結(jié)合。14.根據(jù)權(quán)利要求1所述的人源5，10-次甲基四氫葉酸合成酶的晶體的K10、R14、R145、D154位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域與ATP結(jié)合。15.根據(jù)權(quán)利要求2所述的人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體中a5的152-161位氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域是與從ATP水解下來(lái)的磷酸基團(tuán)結(jié)合的結(jié)構(gòu)域。16.根據(jù)權(quán)利要求4所述的人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體中|33和P4之間的氨基酸即第89和134位氨基酸之間的氨基酸在空間中所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域形成一個(gè)loop環(huán)結(jié)構(gòu)，該結(jié)構(gòu)中的氨基酸與谷氨酸結(jié)合。17.根據(jù)權(quán)利要求5所述的人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體中氨基酸殘基F55、L56、S57、M58、E61、P81、Y83、M90、W109、P135、R148、K150、Y152和Y153所形成的結(jié)構(gòu)域，其特征在于所述結(jié)構(gòu)域形成一個(gè)球形口袋，N5-亞胺磷酸過(guò)渡態(tài)的蝶呤環(huán)位于所述球形口袋的中間。18.根據(jù)權(quán)利要求5所述的人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸過(guò)渡態(tài)的復(fù)合物的晶體中蝶呤環(huán)在空間中的結(jié)構(gòu)域，其特征在于所述蝶呤環(huán)以C4a與C8a之間的鍵為軸發(fā)生了扭轉(zhuǎn)，所述喋呤環(huán)上的電子發(fā)生了重排。19.根據(jù)權(quán)利要求6所述的人源5，10-次甲基四氫葉酸合成酶與10-甲酰四氫葉酸的復(fù)合物的晶體中Y83在空間中的結(jié)構(gòu)域，其特征在于Y83的位置與在權(quán)利要求5所述的晶體的位置相比，發(fā)生了變化。20.與權(quán)利要求12-19中任一項(xiàng)所述的結(jié)構(gòu)域類(lèi)似或者與所述結(jié)構(gòu)域結(jié)合的小分子化合物、短肽、多肽、核酸、抗體或免疫結(jié)合物中的一種或幾種的組合物。全文摘要本發(fā)明涉及一種人源5，10-次甲基四氫葉酸合成酶(MTHFS)的晶體及其結(jié)晶的方法，同時(shí)也涉及人源5，10-次甲基四氫葉酸合成酶與ADP的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與谷氨酸的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與N5-亞胺磷酸的復(fù)合物的晶體及其結(jié)晶的方法、人源5，10-次甲基四氫葉酸合成酶與產(chǎn)物10-甲酰四氫葉酸復(fù)合物的晶體及其結(jié)晶的方法；還涉及這些晶體中重要的結(jié)構(gòu)域或肽片段；以及利用這些結(jié)構(gòu)域或肽片段進(jìn)行藥物設(shè)計(jì)和篩選的方法。文檔編號(hào)C07K16/40GK101781637SQ20091013447公開(kāi)日2010年7月21日申請(qǐng)日期2009年4月17日優(yōu)先權(quán)日2009年4月17日發(fā)明者劉志杰,宋高潔,尼爾·肖恩,成沖云,李楊,武棟申請(qǐng)人:中國(guó)科學(xué)院生物物理研究所