本專利申請要求2014年12月10日提交的名稱為“基因修飾苯丙酮酸脫羧酶、其制備方法和其用途”的美國專利申請第62/089,912號的權(quán)益，所述專利申請以全文引用的方式并入本文中。本發(fā)明涉及在微生物中使用生物酶產(chǎn)生c6-c10化合物，如醇、羧酸和烷烴的領(lǐng)域。更確切地說，其涉及使用一或多種工程改造硫胺素依賴性脫羧酶轉(zhuǎn)化給定2-酮酸底物的領(lǐng)域。表達(dá)如下文所描述的表示m461v變異體的本發(fā)明的基因修飾脫羧酶的一個(gè)特定實(shí)施例的微生物樣品已于2015年12月9日寄存于美國組織類型收集(atcc)專利庫,10801universityblvd.,manassas,va20110。地理政治和環(huán)境問題已激勵(lì)全世界的研究人員追求使用可再生途徑，包括(但不限于)使用微生物的發(fā)酵產(chǎn)生基于石化品的產(chǎn)品。但是，由于微生物通常未能在經(jīng)濟(jì)上可行的速率或產(chǎn)率下產(chǎn)生許多基于石化品的產(chǎn)品，代謝工程改造已廣泛用于建立路徑和/或朝向所關(guān)注的路徑引導(dǎo)代謝物。當(dāng)前，乙醇為使用微生物制得的最常見生物化學(xué)物質(zhì)。但是，在生物燃料和化學(xué)行業(yè)中積極地追求產(chǎn)生較長鏈醇和羧酸的經(jīng)濟(jì)上可行的方法。通過微生物發(fā)酵產(chǎn)生天然氨基酸中的成功尤其在使用氨基酸生物合成路徑產(chǎn)生所關(guān)注的化學(xué)品，包括較長鏈醇和羧酸中產(chǎn)生重大興趣。參見例如becker,j.；wittmann,c.“用于氨基酸生產(chǎn)的系統(tǒng)和合成代謝工程改造-工業(yè)菌株發(fā)展的脈搏(systemsandsyntheticmetabolicengineeringforaminoacidproduction-theheartbeatofindustrialstraindevelopment),”《生物技術(shù)當(dāng)前觀點(diǎn)(curr.opinbiotechnol)》.,2012,23:718-726；和becker,j.；wittmann,c.“基于生物的化學(xué)品、材料和燃料的生產(chǎn)-谷氨酸棒狀桿菌作為通用細(xì)胞工廠(bio-basedproductionofchemicals,materialsandfuels-corynebacteriumglutamicumasversatilecellfactory),”《現(xiàn)代生物技術(shù)觀點(diǎn)(curr.opin.biotechnol.)》,2012,23:631-640。在氨基酸生物合成期間為關(guān)鍵中間物的2-酮酸能經(jīng)受可用于細(xì)胞內(nèi)部的化學(xué)物質(zhì)的生物合成的不同類型的修飾。參見例如gronenberg,l.s.；marcheschi,r.j.；liao,j.c.“原核生物中的下一代生物燃料工程改造(nextgenerationbiofuelengineeringinprokaryotes),”《化學(xué)生物學(xué)新見(curr.opin.chem.biol.)》,2013,17:462-471。在一個(gè)實(shí)例中，美國專利8,232,089描述表達(dá)產(chǎn)生異丁醇的代謝路徑的重組酵母菌，所述路徑包括在與異丁醇產(chǎn)生基因共表達(dá)時(shí)將2-酮基異戊酸酯轉(zhuǎn)化為異丁醛的巴西固氮螺菌脫羧酶。在另一實(shí)例中，美國專利8,298,798描述通過2-酮酸的脫羧，接著通過乳酸乳桿菌(l.lactis)酮基異戊酸酯脫羧酶和酵母菌醇脫氫酶adh6的表達(dá)來還原產(chǎn)生的醛而在大腸桿菌(e.coli)細(xì)胞中產(chǎn)生直鏈和分支鏈醇。也參見atsumi,s.；hanai,t.；liao,j.c.“用于合成分支鏈高級醇作為生物燃料的非發(fā)酵路徑(non-fermentativepathwaysforsynthesisofbranched-chainhigheralcoholsasbiofuels)”《自然(nature)》,2008,451:86-89；marcheschi,r.j.；li,h.；zhang,k.；noey,e.l.；kim,s.；chaubey,a.；houk,k.n.；liao,j.c.“用于碳鏈延長的合成遞歸“+1”路徑(syntheticrecursive“+1”pathwayforcarbonchainelongation),”《acs化學(xué)生物學(xué)(acschem.biol)》.,2012,7:689-697；和zhang,k.；sawaya,m.r.；eisenberg,d.s.；liao,j.c.“用于生物合成非天然醇的擴(kuò)張代謝(expandingmetabolismforbiosynthesisofnonnaturalalcohols),”《美國國家科學(xué)院院刊(proc.natl.acad.sci.u.s.a.)》,2008,105:20653-20658。已經(jīng)由脫羧酶和醛去氫酶的表達(dá)展示2-酮酸中間物在細(xì)胞內(nèi)部轉(zhuǎn)化為羧酸。參見例如xiong,m.；deng,j.；woodruff,a.p.；zhu,m.；zhou,j.；park,s.w.；li,h.；yao,f.“脂族酮的生物催化方法(abio-catalyticapproachtoaliphaticketones),”2012,《科學(xué)報(bào)導(dǎo)(sci.rep.)》2:311；和zhang,k.；woodruff,a.p.；xiong,m.；zhou,j.；dhande,.,k.“用于生產(chǎn)平臺化學(xué)異丁酸的合成代謝路徑(asyntheticmetabolicpathwayforproductionoftheplatformchemicalisobutyricacid),”《化學(xué)與可持續(xù)性、能源與材料(chemsuschem)》,2011,4:1068-1070。經(jīng)由大腸桿菌的leua基因產(chǎn)物的工程改造延長細(xì)胞內(nèi)部的2-酮酸的長度的可行性也已擴(kuò)展可產(chǎn)生自2-酮酸的生物化學(xué)物質(zhì)的范圍。參見例如atsumi,s.,同上.,和zhang,k.,同上。在大腸桿菌中，leuabcd基因延長2-酮酸的長度一個(gè)碳單元，如在亮氨酸生物合成期間所觀測，其中其共同起作用以將2-酮基異戊酸酯(5碳酸)轉(zhuǎn)化為2-酮基異己酸酯(6碳酸)。marcheschi等人.《acs化學(xué)生物學(xué)》.,2012,7:689-697描述leua的活性位點(diǎn)的擴(kuò)展和c4酮酸、2-酮基丁酸酸[2-酮基丁酸酯]延長為c9酮酸、2-酮基壬酸[2-酮基壬酸酯]。盡管有可能使用代謝工程改造在微生物中產(chǎn)生變化長度的醇和羧酸，但到目前為止尚未展示特定c6-c8醇或酸的產(chǎn)生，優(yōu)選地以按總醇產(chǎn)物計(jì)的大于20重量％(wt％)、更優(yōu)選地大于30wt％的量。若干因素似乎決定產(chǎn)生自細(xì)胞內(nèi)部的2-酮酸的醇/酸的特異性。脫羧酶接受變化長度的2-酮酸的雜亂性導(dǎo)致變化長度的醛，其接著通過對應(yīng)的共表達(dá)醛或醇脫氫酶氧化或還原。因此，較高水準(zhǔn)的雜亂性，即較低水準(zhǔn)的特異性導(dǎo)致較高數(shù)量的產(chǎn)物。這轉(zhuǎn)而可意味著尤其所需的特定產(chǎn)物的較低產(chǎn)率。經(jīng)由2-酮酸特定產(chǎn)生醇或羧酸也可受就leuabcd來說的脫羧酶的表達(dá)量不利地影響。較高水準(zhǔn)的具有廣泛底物特異性的脫羧酶傾向于與leua基因產(chǎn)物競爭2-酮酸中間物且進(jìn)而限制路徑延長2-酮酸的能力。這可導(dǎo)致形成比可能需要短的醇或羧酸，再次導(dǎo)致非所需產(chǎn)物和/或產(chǎn)物混合物。一般來說，改良工業(yè)微生物的方法從傳統(tǒng)菌株改良(csi)的隨機(jī)方法到代謝工程改造的高度合理方法變動。csi一般對于緩解產(chǎn)物抑制或提高產(chǎn)率有效，但在產(chǎn)生能夠產(chǎn)生完全新穎產(chǎn)物的菌株中效果小得多。此外，csi為時(shí)間和資源密集型的。為了獲得對于抑制性發(fā)酵產(chǎn)物具有高耐受性的菌株，有必要通過在增加的抑制劑濃度存在下在培養(yǎng)基中連續(xù)培養(yǎng)菌株而連續(xù)篩選和選擇突變體。這通常與使用化學(xué)誘變劑和/或紫外線(uv)輻射的誘導(dǎo)性突變誘發(fā)結(jié)合進(jìn)行。但是，常規(guī)的培養(yǎng)物篩選過程是冗長、耗時(shí)并且通常徒勞的。代謝修飾一般在創(chuàng)造產(chǎn)生新產(chǎn)物的菌株中更有效。這是因?yàn)榛?，且在一些情況下甚至整個(gè)路徑可在生物體之間轉(zhuǎn)移(重組方法)和/或酶可經(jīng)修飾(工程改造方法)。這些方法避免csi的一些缺點(diǎn)。包含重組和工程改造方法兩者的術(shù)語代謝工程改造為廣泛用于設(shè)計(jì)菌株以通過代謝通量分布的改變而實(shí)現(xiàn)代謝物過度產(chǎn)生中的較高效率的定向且通常較快的方法。到目前為止此操作的大部分是關(guān)于使用具有充分研究的遺傳學(xué)和生理學(xué)的生物體(例如大腸桿菌、酵母菌和融合瘤細(xì)胞)來產(chǎn)生次級代謝物(如抗生素)、氨基酸(例如賴氨酸)和異源蛋白質(zhì)。代謝通量分布的化學(xué)計(jì)量分析提供適當(dāng)代謝修飾、最優(yōu)培養(yǎng)基調(diào)配和饋入策略和生物工藝優(yōu)化的指導(dǎo)。但是，此方法仍需要發(fā)酵細(xì)胞中的代謝和調(diào)節(jié)網(wǎng)絡(luò)的深入知識。盡管這些合理方法已在涉及單個(gè)基因或單一基因簇內(nèi)的若干基因的情況下取得成功，但其通常在涉及更復(fù)雜或在很大程度上未知的代謝途徑的情況下無效。這是因?yàn)檫@通常每次靶向一個(gè)基因，且因此未能預(yù)測給定路徑中的多個(gè)基因之間的復(fù)雜相互作用。通過修飾基因密碼的所述部分，即生物體dna進(jìn)行酶修飾，其對應(yīng)于酶的表達(dá)。酶修飾可產(chǎn)生完全新穎的官能團(tuán)且可用于提高所需中間物或產(chǎn)物的特異性或效率。另外，已知某些酶為雜亂的且可發(fā)現(xiàn)進(jìn)行超出其已知自然角色的任務(wù)。此類酶也可經(jīng)修飾以進(jìn)行新轉(zhuǎn)化，但到目前為止，此方法的成功頻繁地限于在商業(yè)上不可行的產(chǎn)率。參見例如zhang,k.,同上。在路徑中修飾多個(gè)酶可理論上用作使特異性和/或催化效率最大化的技術(shù)。已知在某些條件下產(chǎn)生辛醇的生物體的一個(gè)實(shí)例為梭菌屬(clostridium)。梭菌屬的各種物種(例如丙酮丁醇梭菌(c.acetobutylicum)、艱難梭菌(c.difficile)和克氏梭菌(c.kluyveri))用于wo2012135731中。公開案描述通過工程改造的梭菌屬物種產(chǎn)生少量正辛醇，連同其它產(chǎn)物，且將對于正辛醇的不佳特異性歸因于生物體表達(dá)或過度表達(dá)β-酮硫解酶(例如bktb)、乙酰基coa乙?；D(zhuǎn)移酶(例如atob)、3-羥基丁?；?coa去氫酶(例如來自梭菌屬的hbd，或paahi)巴豆酸酶(例如crt)和反烯?；?coa還原酶(例如ter)的能力。一般來說，工程改造修飾是針對生物體的coa路徑以產(chǎn)生高級醇，且此路徑避免發(fā)現(xiàn)于許多梭菌屬物種中的涉及氧敏感性酶和中間物的丁醇產(chǎn)生路徑。顯示為已經(jīng)由本發(fā)明產(chǎn)生的正辛醇的量太小而在商業(yè)上不可行。也參見例如lee,j.y.；jang,y.s.；lee,j.；papoutsakis,e.t.；lee,s.y.“丙酮丁醇梭桿菌m5的代謝工程改造以用于高度選擇性丁醇生產(chǎn)(metabolicengineeringofclostridiumacetobutylicumm5forhighlyselectivebutanolproduction),”《生物技術(shù)學(xué)刊(biotechnol.)》2009,4:1432-1440；和wang,y.；blaschek,h.p.“通過用拜氏梭菌發(fā)酵而優(yōu)化從熱帶玉米稈汁的丁醇生產(chǎn)(optimizationofbutanolproductionfromtropicalmaizestalkjuicebyfermentationwithclostridiumbeijerinckii),”《生物資源技術(shù)(bioresour.technol.)》,2011,102:9985-9990。當(dāng)前研究的基因工程改造的一種應(yīng)用是在能源領(lǐng)域。關(guān)于未來獲得和使用化石燃料的稀缺、成本和環(huán)境影響的擔(dān)憂已刺激在采用廉價(jià)、可再生生物質(zhì)作為由其制得的燃料和化學(xué)品兩者的替代來源中的興趣。由于原油價(jià)格變得更易變，基于生物的化學(xué)品和工業(yè)產(chǎn)品已變?yōu)槠涫脱苌詫?yīng)物的有吸引力的替代方案。使用厭氧微生物的發(fā)酵方法提供將生物質(zhì)和農(nóng)業(yè)廢棄物轉(zhuǎn)化為有用產(chǎn)物的有前景的途徑，且同時(shí)整改可在處置低價(jià)值農(nóng)業(yè)商品和食品加工副產(chǎn)物/廢棄物中遇到的問題?？捎傻统杀旧镔|(zhì)進(jìn)料制備的有用產(chǎn)物中的一些為有機(jī)酸和醇，包括辛醇。c6-c10醇尤其用作制備作為許多行業(yè)中的高度所需進(jìn)料化學(xué)品的烷烴、烯烴和醛的低成本起始物質(zhì)。這些行業(yè)包括用作溶液聚合的共單體，和洗滌劑行業(yè)，其使用烷基化酚的這些前驅(qū)物來生產(chǎn)洗滌劑前驅(qū)物。這些醇也可用作表面活性劑；潤膚劑；化妝品和食品行業(yè)種的增稠劑；殺蟲劑；和多種其它應(yīng)用。在一個(gè)實(shí)施例中，本發(fā)明提供一種基因修飾微生物的方法，其包含(a)選擇產(chǎn)生c7-c112-酮酸的微生物；和(b)插入編碼對應(yīng)于seqid4、8、14、16、18、28、30、32、34、36、38、40、42、46、52、54、56、62、64、66、68或76的氨基酸序列，或與其至少90％同源的氨基酸序列的非天然核酸序列；使得非天然苯丙酮酸脫羧酶表達(dá)于微生物中。在另一實(shí)施例中，本發(fā)明提供基因修飾微生物。在另一實(shí)施例中，本發(fā)明提供一種制備c6-c10醛、c6-c10醇、c6-c10羧酸或c6-c10烷烴的方法，其包含以下步驟：(a)使2-酮基丁酸酯或2-酮基異戊酸酯、異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶和異丙基蘋果酸去氫酶在使得2-酮基丁酸酯或2-酮基異戊酸酯轉(zhuǎn)化為c7-c112-酮酸的條件下接觸；(b)使c7-c112-酮酸與苯丙酮酸脫羧酶在使得c7-c112-酮酸轉(zhuǎn)化為碳原子比轉(zhuǎn)化的c7-c112-酮酸少一個(gè)的c6-c10醛的條件下接觸，所述苯丙酮酸脫羧酶由編碼對應(yīng)于seqid4、8、14、16、18、28、30、32、34、36、38、40、42、46、52、54、56、62、64、66、68或76的氨基酸序列，或與其至少90％同源的氨基酸序列的非天然核酸序列表示；和(c)任選地使c6-c10醛與(1)醇脫氫酶在形成c6-c10醇的條件下接觸；或(2)醛去氫酶在形成c6-c10羧酸的條件下接觸；或(3)脂肪醛脫羧酶在形成c6-c10烷烴的條件下接觸；進(jìn)行所述方法以使得在微生物內(nèi)部或外部和好氧或厭氧條件下獨(dú)立地進(jìn)行每一步驟和子步驟。在另一實(shí)施例中，本發(fā)明提供一種多肽，其包含對應(yīng)于seqid8、14、16、18、28、30、32、34、36、38、40、42、46、52、54、56、62、64、66、68或76，或與其至少90％同源的氨基酸序列，主要由所述氨基酸序列組成，或由所述氨基酸序列組成。在另一實(shí)施例中，本發(fā)明提供一種基因修飾微生物，其包含(a)2-酮酸源；(b)將2-酮酸轉(zhuǎn)化為c7-c11醛的野生型代謝路徑；和(c)由編碼與基因庫寄存編號l26240對應(yīng)的氨基酸序列，或與其至少80％同源的氨基酸序列的核酸序列表示的非天然苯丙酮酸脫羧酶；此類序列任選地經(jīng)以下各者修飾：(1)經(jīng)纈氨酸取代met-380；或(2)經(jīng)纈氨酸、亮氨酸或異亮氨酸取代phe-385；或(3)經(jīng)纈氨酸、亮氨酸、丙氨酸或半胱氨酸取代met-461；或(4)經(jīng)纈氨酸或亮氨酸取代phe-465；或(5)經(jīng)甘氨酸、丙氨酸、纈氨酸或亮氨酸取代phe-532；或(6)經(jīng)纈氨酸、亮氨酸、異亮氨酸、丙氨酸或甘氨酸取代gln-536；或(7)如(1)-(6)中所述的兩個(gè)或三個(gè)取代的任何組合對于下文描述的序列，每一奇數(shù)編號的序列標(biāo)識號(seqid)顯示核苷酸或核酸序列，且每一偶數(shù)編號的seqid顯示對應(yīng)編碼的氨基酸序列。核酸序列編碼氨基酸序列，其中術(shù)語“氨基酸序列”等效于“多肽”或“蛋白質(zhì)”。所有提到的蛋白質(zhì)(偶數(shù)編號的)seqid承認(rèn)有可能使用替代密碼子產(chǎn)生給定氨基酸序列的事實(shí)。seqid1和2表示附接于氨基酸序列的起點(diǎn)的13個(gè)氨基酸的組氨酸標(biāo)簽。seqid3和4表示對應(yīng)于基因庫寄存編號l26240的野生型巴西固氮螺菌(azospirillumbrasilense)苯丙酮酸脫羧酶基因。seqid5和6表示seqid3和4的基因序列，但abppdc中的met-380在位置380處經(jīng)纈氨酸置換(基于無his-tag的氨基酸序列；在13個(gè)氨基酸的his-tag的情況下，這將為位置393)。修飾帶有名稱m380v，因此遵守行業(yè)標(biāo)準(zhǔn)，其中氨基酸修飾定義為原始單字母氨基酸密碼，接著為氨基酸位置，接著為新氨基酸單字母密碼。seqid7和8表示f385l。seqid9和10表示f385v。seqid11和12表示f385i。seqid13和14表示m461c。seqid15和16表示m461v。seqid17和18表示m461l。seqid19和20表示m461a。seqid21和22表示f465l。seqid23和24表示f532a。seqid25和26表示f532g。seqid27和28表示f532v。seqid29和30表示f532l。seqid31和32表示q536g。seqid33和34表示q536a。seqid35和36表示q536l。seqid37和38表示q536i。seqid39和40表示q536v。seqid41和42表示f532v/q536v。seqid43和44表示m380l/m461v。seqid45和46表示m380v/m461v。seqid47和48表示f385v/m461v。seqid49和50表示f385l/m461v。seqid51和52表示f532a/q536v。seqid53和54表示f532v/q536a。seqid55和56表示f385l/q536v。seqid57和58表示f385v/q536v。seqid59和60表示m461v/q536v。seqid61和62表示m461l/q536v。seqid63和64表示m461a/q536v。seqid65和66表示m461v/f532v。seqid67和68表示f465l/q536v。seqid69和70表示f465v/q536v。seqid71和72表示f465l/f532v。seqid73和74表示f532a/q536a。seqid75和76表示m461v/f532v/q536v。seqid77和78表示m380v/m461v/q536v。seqid79和80表示f385l/m461l/q536v。seqid81和82表示m380v/f385v/m461v。圖1說明通過遞歸(迭代)活性，接著脫羧為比迭代路徑中在其之前的2-酮酸短一個(gè)碳原子的醛的鏈延長。圖2說明使用異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶、異丙基蘋果酸去氫酶和醇脫氫酶(adh6)以及abppdc的f385l變異體(seqid8)的組合從2-酮基丁酸酯體外產(chǎn)生直鏈c5-c8醇。圖3說明使用異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶、異丙基蘋果酸去氫酶和醇脫氫酶(adh6)以及abppdc的f385l變異體(seqid8)的組合從2-酮基異戊酸酯體外產(chǎn)生分支鏈c5-c8醇。圖4說明使用異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶、異丙基蘋果酸去氫酶和醇脫氫酶(adh6)以及abppdc的f385l變異體(seqid8)的組合從3-甲基-2-酮基戊酸酯體外產(chǎn)生分支鏈c5-c8醇。圖5說明含有“+1路徑”酶以及abppdc野生型(wt)、abppdc變異體、kivdwt(雷特氏乳球菌酮基異戊酸酯脫羧酶)且不含脫羧酶的大腸桿菌的血清瓶發(fā)酵的平均醇分布。adh6也包含于所有菌株構(gòu)筑體中。一般來說，本發(fā)明尤其包括新苯丙酮酸脫羧酶的兩個(gè)特定實(shí)施例，其可在第一實(shí)施例中為對應(yīng)于基因庫寄存編號l26240的獲自巴西固氮螺菌(a.brasilense)或與其至少80百分比(％)同源的氨基酸序列的表達(dá)。在第二實(shí)施例中，本發(fā)明包括先前定義的基因修飾苯丙酮酸脫羧酶，但具有另外的有意基因工程改造以在序列內(nèi)插入特定氨基酸的一個(gè)、兩個(gè)或三個(gè)修飾，其同樣用于以在許多情況下在進(jìn)行多種生物合成(其中苯丙酮酸脫羧酶參與)中有利的方式修改苯丙酮酸脫羧酶的催化效率。確切地說，野生型或核酸修飾的巴西固氮螺菌苯丙酮酸脫羧酶可與異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶和異丙基蘋果酸去氫酶組合以產(chǎn)生醇、羧酸或烷烴。如將在本文中顯示，相比于所選宿主微生物的野生型酶具有改進(jìn)的特性的新苯丙酮酸脫羧酶是通過以描述于本文中的多種方式中的一種進(jìn)行基因修飾而產(chǎn)生；或?yàn)橛砂臀鞴痰菥奖崦擊让钢辽?0％同源且包括相同修飾的氨基酸序列表示的酶；經(jīng)由重組、工程改造或組合重組和工程改造方法的技術(shù)制造其的方法；使用野生型或新苯丙酮酸脫羧酶制造c6-c10醇、羧酸和烷烴的方法；以及可表達(dá)或過度表達(dá)此酶且可用于產(chǎn)生c6-c10醇、羧酸和烷烴的基因修飾微生物有機(jī)體。在本文使用所述術(shù)語時(shí)，同源性是指序列中列出的氨基酸在其給定位置的80％或更大的一致或功能對應(yīng)。新苯丙酮酸脫羧酶可在某些特定實(shí)施例中用作或表示為以下代謝路徑的一部分：經(jīng)由合成代謝(例如伍德-永達(dá)爾(wood-ljundahl))或分解代謝(例如糖酵解或磷酸戊糖路徑)途徑產(chǎn)生乙?；鵦o-a，且最終參與c7-c112-酮酸的轉(zhuǎn)化以形成少一個(gè)碳的對應(yīng)c6-10醛。在一些實(shí)施例中，c6-c10醛可進(jìn)一步反應(yīng)以形成c6-c10醇、羧酸或烷烴。本發(fā)明的基因修飾苯丙酮酸脫羧酶由于描述于本文中的其氨基酸序列中的特異性改變而提供對各種底物的特異性中的一些顯著差異，且特異性的此改變提供就產(chǎn)物產(chǎn)率和減少或消除非所需和/或競爭副產(chǎn)物來說的重要優(yōu)點(diǎn)。本發(fā)明包括巴西固氮螺菌苯丙酮酸脫羧酶的多種改變的氨基酸序列，其已鑒別為相比于seqid4中顯示的對應(yīng)于基因庫寄存編號l26240的野生型巴西固氮螺菌氨基酸序列展現(xiàn)改進(jìn)的c7-c112-酮酸的脫羧。野生型序列內(nèi)的六個(gè)位點(diǎn)已鑒別為對于獲得所述改進(jìn)至關(guān)重要。這些為met-380、phe-385、met-461、phe-465、phe-532、gln-536和其組合。在每一改變中作出變化，其中纈氨酸、亮氨酸、丙氨酸、甘氨酸或異亮氨酸在野生型氨基酸的鑒別位點(diǎn)處經(jīng)取代，其中取代從單位點(diǎn)(即，構(gòu)成三個(gè)堿基對單一氨基酸)取代到多種定義為優(yōu)選地2到3個(gè)鑒別位點(diǎn)的“組合”的多位點(diǎn)(2到5個(gè)位點(diǎn))取代改變。seqid3-82顯示包括指定的取代中的一或多者的產(chǎn)生的許多變化形式的氨基酸序列。取代可概述如下：(1)用纈氨酸取代met-380；或(2)用纈氨酸、亮氨酸或異亮氨酸取代phe-385；或(3)用纈氨酸、亮氨酸、丙氨酸或半胱氨酸取代met-461；或(4)用纈氨酸或亮氨酸取代phe-465；或(5)用甘氨酸、丙氨酸、纈氨酸或亮氨酸取代phe-532；或(6)用纈氨酸、亮氨酸、異亮氨酸、丙氨酸或甘氨酸取代gln-536；或(7)如(1)-(6)中所述的三種取代的任何組合；所屬領(lǐng)域的技術(shù)人員應(yīng)了解，本發(fā)明的基因修飾苯丙酮酸脫羧酶可在體內(nèi)(即，通過基因修飾微生物)或體外使用。鑒于此，如本文所用的術(shù)語“經(jīng)基因修飾”或“經(jīng)修飾”是指具有刻意改變的氨基酸序列，即“非野生型”氨基酸序列的本發(fā)明苯丙酮酸脫羧酶的群組，或具有關(guān)于(至少)在本文中描述和定義為發(fā)明性的特定經(jīng)修飾脫羧酶已刻意改變的基因組的微生物(取決于作為形容詞的任一術(shù)語的放置)。此類改變可經(jīng)由重組技術(shù)實(shí)現(xiàn)，其中一或多種基因從第二不同微生物轉(zhuǎn)移至目標(biāo)微生物中；或工程改造技術(shù)，其中目標(biāo)微生物內(nèi)的核酸一般經(jīng)由定點(diǎn)突變誘發(fā)改變，導(dǎo)致至少一種核酸轉(zhuǎn)化為不同核酸并且因此修飾一或多種酶。在當(dāng)今dna合成技術(shù)的情況下，重組技術(shù)也可使用完全合成dna實(shí)現(xiàn)，所述dna使用常規(guī)方法轉(zhuǎn)移至目標(biāo)微生物。也可采用以上方法中的任一種的組合。本發(fā)明進(jìn)一步包括一種制備c6-c10醛、c6-c10羧酸、c6-c10烷烴和c6-10醇(如己醇、庚醇和/或1-辛醇)的方法，其經(jīng)由起始底物與包括本發(fā)明的基因修飾苯丙酮酸脫羧酶中的一或多種的一系列酶之間的接觸以使用另外的酶和步驟將底物最終轉(zhuǎn)化為所需c6-c10醛、醇、羧酸或烷烴。此方法可在非天然存在，即基因工程改造的細(xì)胞的所描述實(shí)施例中的一個(gè)中，即在非天然存在的微生物中以生物合成方式進(jìn)行；或c6-c10醇、羧酸或烷烴的生產(chǎn)可經(jīng)由體外方法進(jìn)行，通常從不包括微生物的起始點(diǎn)開始。為了獲得本發(fā)明的經(jīng)修飾苯丙酮酸脫羧酶的群組，在一個(gè)實(shí)施例中需要與下文所描述類似地執(zhí)行方案。一般來說，實(shí)例顯示涉及改變給定密碼子中的一或多個(gè)核酸堿基以改變核酸堿基為一部分的酶的工程改造的基因修飾。這可僅用于產(chǎn)生用于例如體外分析目的的改變的酶。相比之下，宿主微生物的基因組可優(yōu)選地經(jīng)改變以用于較大規(guī)模生產(chǎn)株?？扇缢鶎兕I(lǐng)域的技術(shù)人員一般理解地進(jìn)行被設(shè)計(jì)用于體外酶生產(chǎn)的以下方法。一般來說，適合的數(shù)據(jù)庫(如基因庫(genbank))用于獲得用于野生型酶的基因編碼，接著識別適合于修飾的密碼子。此識別可用作蛋白質(zhì)工程的技術(shù)已知方法的基礎(chǔ)，其中計(jì)算機(jī)分子建模鑒別可有效地采用酶/底物界面的修飾的結(jié)構(gòu)位置并且使得能夠區(qū)別所述結(jié)構(gòu)位置。接著使用分子生物學(xué)技術(shù)進(jìn)行給定所需修飾，其中經(jīng)由定點(diǎn)突變誘發(fā)進(jìn)行一或多個(gè)核酸堿基的改變。變異型酶必須接著經(jīng)受純化以分離出非目標(biāo)蛋白質(zhì)，留下將展現(xiàn)高于野生型的催化效率的純化酶。這可根據(jù)最適合于給定特定酶的方法經(jīng)恰當(dāng)?shù)伢w外分析。顯示具有所需催化效率水準(zhǔn)的分析的酶進(jìn)而確認(rèn)為所需基因修飾的產(chǎn)物，且可用于體外生產(chǎn)方法，如將c7-c112-酮酸體外轉(zhuǎn)化為少一個(gè)碳的對應(yīng)c6-c10醛(例如將2-酮基壬酸酯轉(zhuǎn)化為辛醛，或?qū)?-酮基辛酸酯轉(zhuǎn)化為庚醛)，其可接著在一個(gè)實(shí)施例中通過與適當(dāng)野生型或非野生型醇脫氫酶接觸而還原，以形成對應(yīng)c6-c10醇。如上文所提到，可在體內(nèi)或體外進(jìn)行本發(fā)明。體內(nèi)方法可對于商業(yè)規(guī)模生產(chǎn)為優(yōu)選的，而在一些情況下，體外方法可適合于商業(yè)規(guī)模生產(chǎn)。通常，體外方法可尤其便于實(shí)驗(yàn)室和一般研究目的，以便進(jìn)行酶分析。舉例來說，可制備適用于酶促產(chǎn)物大規(guī)?；蛏虡I(yè)規(guī)模發(fā)酵生產(chǎn)，如在某些特定實(shí)施例中，c6-c10醇或c6-c10醇的組合的所需微生物有機(jī)體。此類制備可如下進(jìn)行：通過使用重組技術(shù)將來自第一微生物的編碼所需改進(jìn)的酶的dna或dna片段插入至第二“宿主”微生物的基因組中，已知或相信所述第二“宿主”微生物具有一或多個(gè)所需代謝路徑和/其它所需特征，如耐抑制發(fā)酵能力。一般來說，體內(nèi)方法采用此類微生物野生型代謝路徑在不同步數(shù)中首先將適合的含碳底物轉(zhuǎn)化為丙酮酸酯，且接著將丙酮酸酯轉(zhuǎn)化為2-酮基丁酸酯，或者，2-酮基異戊酸酯。舉例來說，在一個(gè)實(shí)施例中，適合的含碳底物，如c5或c6糖(例如葡萄糖、蔗糖、戊糖或其組合)可經(jīng)由分解代謝或合成代謝路徑中的一個(gè)，如糖酵解或磷酸戊糖路徑直接轉(zhuǎn)化為丙酮酸鹽。此后，丙酮酸鹽可經(jīng)由pc(丙酮酸羧化酶)；aat(天冬氨酸轉(zhuǎn)氨酶)；thrabc(thra，其為雙官能天冬氨酸激酶/高絲氨酸去氫酶；thrb，其為高絲氨酸激酶；thrc，其為蘇氨酸合成酶；且asd，其為天冬氨酸半醛去氫酶)首先轉(zhuǎn)化為l-蘇氨酸。l-蘇氨酸接著經(jīng)由ilva(蘇氨酸脫水酶)轉(zhuǎn)化為2-酮基丁酸酯。在一替代實(shí)施例中，丙酮酸鹽可經(jīng)由亮氨酸生物合成中的ilvbn/ilvgm、ilvc和ilvd的活性轉(zhuǎn)化為2-酮基異戊酸酯。另外參見zhang,k.；sawaya,m.r.等人,同上。從這個(gè)角度，涉及拉長2-酮酸的亮氨酸生物合成路徑內(nèi)的野生型或基因修飾形式的三種酶中的一或多者經(jīng)操作以將2-酮基丁酸酯或2-酮基異戊酸酯轉(zhuǎn)化為c7-c112-酮酸。在不提及任何特定微生物的情況下，這些酶一般稱為異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶和異丙基蘋果酸去氫酶。但是，特定在大腸桿菌中，其分別稱為leua(基因庫寄存編號nc000913.3，基因標(biāo)識：947465)、leub(基因庫寄存編號nc000913.3，基因標(biāo)識：944798)和leucd(基因庫寄存編號nc000913.3，基因標(biāo)識945076和基因標(biāo)識：945642)。此鏈延長的一個(gè)實(shí)例顯示于圖1中，其中2-酮基丁酸酯經(jīng)由稱為“+1路徑”的多個(gè)步驟轉(zhuǎn)化為c7-c112-酮酸2-酮基壬酸酯。在某些特定實(shí)施例中，涉及延伸2-酮酸的的亮氨酸生物合成路徑的野生型酶可尤其通過包括至少一種外源酶、酶復(fù)合物或其組合而修飾以在發(fā)生鏈延長時(shí)將2-酮基丁酸酯首先轉(zhuǎn)化為2-酮基戊酸酯，接著轉(zhuǎn)化為2-酮基己酸酯，接著轉(zhuǎn)化為2-酮基庚酸酯且必要時(shí)繼續(xù)轉(zhuǎn)化為另一延長的2-酮酸直至轉(zhuǎn)化為2-酮基十一酸酯，即所需c7-c112-酮酸。但是，任選地可能僅修飾酶、酶復(fù)合物或其組合中的一個(gè)或兩個(gè)以便獲得可接受或所需產(chǎn)量的c7-c112-酮酸。這些酶可包括如上文所提及的leua、leub和/或leucd。尤其適用于路徑的此部分的修飾的為2014年12月10日提交的同在申請中的國際專利申請序號pct/us14/69438(代理人案號75413-wo-pct)的公開內(nèi)容，所述專利申請要求2013年12月12日提交的美國臨時(shí)專利申請第61/915,040號(代理人案號75413-us-psp)的權(quán)益，其均以全文引用的方式并入本文中。在某些實(shí)施例中，至少選擇經(jīng)修飾的異丙基蘋果酸去氫酶變異體(其為大腸桿菌中的leub基因的產(chǎn)物)，或在其它實(shí)施例中，至少包括經(jīng)修飾的leua(leua')和leub'變異體，其優(yōu)選地但未必如引用的專利申請中的一個(gè)或兩個(gè)中所述。也優(yōu)選地采用leua'、leub'和經(jīng)修飾的leucd(leucd')酶/酶復(fù)合物的其它組合。同樣，應(yīng)注意“l(fā)eu”+字母(a、b、cd)名稱為大腸桿菌中的異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶和異丙基蘋果酸去氫酶的亮氨酸路徑酶的特定名稱，而其它生物體的亮氨酸路徑中的相同或等效酶可具有不同名稱。最后，本發(fā)明的基因修飾苯丙酮酸脫羧酶可在此特定實(shí)施例中用以將c7-c112-酮酸轉(zhuǎn)化為比底物2-酮酸少一個(gè)碳的醛。在各種實(shí)施例中，所得c6-c10醛可發(fā)現(xiàn)多種用途，作為產(chǎn)物本身或作為用于生產(chǎn)包括c6-c10醇的產(chǎn)物的起始或中間產(chǎn)物。可經(jīng)由c6-c10醛通過適當(dāng)野生型或基因修飾醇脫氫酶的轉(zhuǎn)化實(shí)現(xiàn)c6-c10醇的制備，但其它產(chǎn)物，如c6-c10烷烴也可經(jīng)由脂肪醛脫羧酶的作用或表達(dá)制備，或c6-10羧酸可通過醛去氫酶的作用或表達(dá)制備。參見例如choi,y.j.；lee,s.y.“短鏈烷烴的微生物生產(chǎn)(microbialproductionofshort-chainalkanes)”,《自然(nature)》,2013,502:571-574。因此，c6-10醛在工業(yè)上高度適用作用于制備多種其它產(chǎn)物的極佳中間產(chǎn)物。因此，預(yù)期本發(fā)明的基因修飾苯丙酮酸脫羧酶家族將可適用于多種行業(yè)。此類行業(yè)可包括(例如)在燃料、塑料、食品、包裝、化妝品、香料、制藥、清潔材料、污染控制、香料、藥物和許多其它行業(yè)中的用途。盡管存在多種完全屬于本發(fā)明的權(quán)利要求的范圍內(nèi)的可能的氨基酸序列，但應(yīng)注意，通過序列標(biāo)識號(seqid)鑒別為選自seqid34、36、38、40、42、46、62、68和76的某些氨基酸序列就對c7-c112-酮酸脫羧基來說是尤其適合和優(yōu)選的。實(shí)例1設(shè)計(jì)對于2-酮基壬酸脫羧具有較高催化效率的巴西固氮螺菌苯丙酮酸脫羧酶(abppdc)變異體abppdc與3-脫氮-硫胺素二磷酸酯和5-苯基-2-氧代戊酸(pdb標(biāo)識碼2q5q)的三元復(fù)合物的晶體結(jié)構(gòu)模型用于鑒別加襯abppdc的活性位點(diǎn)內(nèi)的2-酮酸結(jié)合袋的殘基。參見例如versees,w.；spaepen,s.；wood,m.d.；leeper,f.j.；vanderleyden,j.；steyaert,j.“二磷酸硫胺素依賴性脫羧酶中的異位底物活化的分子機(jī)制(molecularmechanismofallostericsubstrateactivationinathiaminediphosphate-dependentdecarboxylase)”,《生物化學(xué)雜志(j.biol.chem.)》,2007,282:35269-35278。將命名為met-380、met-461、phe-385、phe-465、gln-536和phe-532的氨基酸位點(diǎn)基于其與5-苯基-2-氧代戊酸的關(guān)系選擇用于取代實(shí)驗(yàn)。如表1中所列地進(jìn)行一或多個(gè)位點(diǎn)的取代且制備變異體。酶f532v以纈氨酸置換abppdc中的phe-532，而酶f532l以亮氨酸置換phe-532。酶f385l/m461v以亮氨酸置換phe-385且以纈氨酸置換met-461。表1中的其余的巴西固氮螺菌苯丙酮酸脫羧酶(abppdc)變異體是根據(jù)氨基酸(第一個(gè)字母，其中“f”表示“苯丙氨酸[phe]”；“m”表示“甲硫氨酸”[met]；且“q”表示“谷氨酰胺”[gln])、其在氨基酸序列中的位置(編號)和用作替換的氨基酸(最后一個(gè)字母，其中“l(fā)”表示“亮氨酸”；“v”表示“纈氨酸”；“a”表示“丙氨酸”；“c”表示“半胱氨酸”；“i”表示“異亮氨酸”；且“g”表示“甘氨酸”)命名。經(jīng)修飾abppdc變異體中的每一個(gè)經(jīng)表達(dá)和純化，且接著測試針對三種底物的活性，所述底物為2-酮基己酸酯(2-kh)、2-酮基辛酸酯(2-ko)和2-酮基壬酸酯(2-kn)。將預(yù)期2-kh、2-ko和2-kn在通過abppdc脫羧時(shí)分別形成戊醛、庚醛和辛醛。使用下文描述的高通量酶分析在兩個(gè)步驟中進(jìn)行abppdc變異體的評估。起初，測試所有變異體針對單一高濃度(2mm)的2-kh和2-kn的活性(如表1中所示)。在初始評估之后，對于選擇數(shù)目的變異體進(jìn)行詳細(xì)動力學(xué)分析以測定最大速率(kcat)、產(chǎn)生半最大速率(k0.5，酶在michaelis-menten動力學(xué)之后的km的等效物)的底物濃度和酶針對2-ko和2-kn的催化效率(kcat/k0.5)(如表2中所示)。對于2-kn具有較高特異性(較高kcat/k0.5)的abppdc變異體將在細(xì)胞內(nèi)部產(chǎn)生衍生自其的辛醛和化學(xué)物質(zhì)中有效。表1.abppdc變異體的序列清單和活性seqid4為巴西固氮螺菌苯丙酮酸脫羧酶(基因庫寄存編號l26240)的氨基酸序列。seqid6-82為設(shè)計(jì)和表示于本發(fā)明中的蛋白質(zhì)的序列。本發(fā)明中表示的所有蛋白質(zhì)在n端具有13個(gè)以組氨酸標(biāo)簽形式添加的額外氨基酸(顯示于seqid2中)。實(shí)例2a.巴西固氮螺菌苯丙酮酸脫羧酶(abppdc)和其工程改造變異體在大腸桿菌中的異源表達(dá)為了評估表1中列出的野生型abppdc和其變異體的底物特異性，所有蛋白質(zhì)的基因分別表達(dá)于大腸桿菌細(xì)胞中且蛋白質(zhì)產(chǎn)物與細(xì)胞分離。從ncbi數(shù)據(jù)庫下載巴西固氮螺菌苯丙酮酸脫羧酶(基因庫寄存編號l26240)的基因序列。將包括六個(gè)(6)組氨酸(his)的13個(gè)額外氨基酸的密碼子添加到abppdc基因序列的met-1密碼子的上游。此類修飾允許具有13個(gè)額外氨基酸的組氨酸標(biāo)記的abppdc在n端上的表達(dá)。額外氨基酸以使用ni-nta色譜在單一步驟中純化蛋白質(zhì)的輔助的形式附接。具有13個(gè)額外氨基酸的整個(gè)abppdc序列(seqid2)經(jīng)化學(xué)合成且接著克隆到syntheticgenomics,inc.(sandiego,ca)的t7聚合酶促進(jìn)劑下游的prsfduet-1載體(emdbiosciences)中。最終載體在裝運(yùn)之前由syntheticgenomics,inc.定序。使用newenglandbiolab的q5定點(diǎn)突變誘發(fā)試劑盒(目錄號e0554s)化學(xué)合成或產(chǎn)生表1中列出的abppdc變異體的基因且克隆至prsfduet-1載體中。含有abppdc或abppdc變異體基因的prsfduet-1載體如下所述地轉(zhuǎn)化成大腸桿菌、abppdc或其變異體，接著表達(dá)且最終純化。接著使用獲自emdbiosciences的感受態(tài)bl21(de3)細(xì)胞進(jìn)行大腸桿菌表達(dá)研究。根據(jù)試劑盒說明書進(jìn)行轉(zhuǎn)化且涉及混合50微升(μl)等分試樣的感受態(tài)細(xì)胞與1μl載體。使用康霉素作為生長培養(yǎng)基中的標(biāo)記物選擇含有abppdc表達(dá)載體的細(xì)胞。接著在含有50微克/毫升(μg/ml)康霉素的盧里亞-貝爾塔尼(luria-bertani；lb)培養(yǎng)液瓊脂板上選擇含有abppdc或abppdc變異體表達(dá)載體的大腸桿菌轉(zhuǎn)化體。在37攝氏度(℃)下培育所述板16小時(shí)(h)。通過將單一群落的轉(zhuǎn)化體轉(zhuǎn)移至50毫升(ml)含有50μg/ml康霉素的lb培養(yǎng)基中且在37℃下在振蕩的情況下以220轉(zhuǎn)/分(rpm)培育過夜而開始起子培養(yǎng)。次日，將7ml起子培養(yǎng)物接種至800mlterrificbroth(tb)中且在37℃下培育培養(yǎng)物直至培養(yǎng)物達(dá)到0.5的600納米下的光學(xué)密度(od600nm)。添加1mm的最終濃度下的異丙基β-d-1-硫代哌喃半乳糖苷(iptg)以誘發(fā)abppdc或abppdc變異體基因的表達(dá)且將培養(yǎng)物轉(zhuǎn)移至15℃恒溫箱后維持16小時(shí)(h)。在16h結(jié)束時(shí)，培養(yǎng)物在8000轉(zhuǎn)/分(rpm)下離心以使細(xì)胞?；⒓?xì)胞團(tuán)分成兩個(gè)等分試樣且儲存于-80℃下過夜，隨后純化。獲自400ml表達(dá)培養(yǎng)物的大腸桿菌細(xì)胞團(tuán)懸浮于含有1μg/mldna酶(thermofisherscientific,inc.,rockford,il)、1μg/ml溶菌酶(thermofisherscientific,inc.,rockford,il)、1毫摩爾(mm)二硫蘇糖醇和蛋白酶抑制劑混合液(rpicorp.,mountprospect,il)的b-per試劑(thermofisherscientific,inc.,rockford,il)中。將懸浮液在室溫下平緩地?fù)u動30分鐘(min)且在15,000倍重力(×g)下離心20min以使細(xì)胞碎片粒化。分離上清液且與5mlco-nta樹脂(thermofisherscientific,inc.,rockford,il)一起培育，所述樹脂已用平衡緩沖液(50mm磷酸鈉，ph8.0，含有300mm氯化鈉、20mm咪唑、50μl蛋白酶抑制劑混合液和15％甘油)預(yù)平衡。在4℃下的1h的培育期之后，用5倍體積的平衡緩沖液洗滌酶結(jié)合樹脂。abppdc或其變異體用過含有200mm咪唑的平衡緩沖液從co-nta樹脂洗脫。洗脫的蛋白質(zhì)相對于磷酸鹽緩沖鹽水滲析且以20％甘油溶液的形式儲存于-20℃下。b.測定abppdc和abppdc變異體的底物特異性使用如在實(shí)例1中詳細(xì)描述的方法進(jìn)行abppdc變異體的底物特異性的評估。進(jìn)行高通量abppdc偶合酶分析以評估abppdc變異體的底物特異性。分析涉及使用醇脫氫酶(adh6，基因庫寄存編號np014051.3)減少產(chǎn)生自abppdc介導(dǎo)的2-酮酸脫羧的醛。abppdc催化反應(yīng)的初始速度測定自在醛的adh6催化的還原期間發(fā)生的還原煙酰胺腺嘌呤二核苷酸磷酸(nadph)的氧化的速率。htp篩選分析涉及用含0.5mm二磷酸硫胺素、0.35mmnadph、4.7微克(μg)酵母菌adh6(基因庫寄存編號np014051.3)和0.3毫克/毫升(mg/ml)牛血清白蛋白(bsa)的abppdc分析緩沖液(50mm3-(n-嗎啉基)丙磺酸，ph6.8，含有2.5mm氯化鎂(mgcl2))在30℃下培育2mm2-kh或2mm2-kn。反應(yīng)通過在30℃下添加含有0.5μg至3.5μg稀釋于含有1mg/mlbsa的abppdc分析緩沖液中的abppdc變異體的工作酶儲備液而開始。含有200μl反應(yīng)混合物的所述板在2500×g下離心15秒且在30℃下預(yù)平衡的biotektm板讀取器上在340nm處以分光光度法追蹤反應(yīng)混合物的吸收率變化。使用340nm處的nadph消耗速率和nadph的消光系數(shù)(6.22mm-1cm-1)計(jì)算酶反應(yīng)的初始速度。所有變異體的活性以存在于反應(yīng)混合物中的酶的量標(biāo)準(zhǔn)化且表示為納摩爾/分鐘/毫克(nmol.min-1.mg-1)。使用購自thermofisherscientific,inc.的來自piercebiotechnologyinc.的660nm總蛋白質(zhì)分析試劑盒，使用bsa作為標(biāo)準(zhǔn)物測定用于使活性標(biāo)準(zhǔn)化的蛋白質(zhì)濃度。2-酮基辛酸酯(2-ko)和2-酮基壬酸酯(2-kn)通過abppdc和其變異體的脫羧的動力學(xué)參數(shù)也使用相同htpabppdc偶合酶分析測定，除了2-ko或2-kn的濃度從0變?yōu)?mm。對于展現(xiàn)底物活化的abppdc變異體，如從相對于底物濃度曲線的初始速度的s形曲線顯而易見，通過使用非線性回歸將數(shù)據(jù)擬合至希爾方程式(hillequation)(表2的圖例中示出)獲得2-酮基酸脫羧的動力學(xué)參數(shù)(kcat、k0.5和kcat/k0.5)。對于遵循正態(tài)飽和動力學(xué)的變異體，通過使用非線性回歸將初始速度擬合至michaelis-menten方程式而動力學(xué)參數(shù)(kcat、km和kcat/km)。使用graphpadprismtm軟件進(jìn)行非線性回歸。表2列出通過abppdc和其變異體的2-ko和2-kn脫羧的動力學(xué)參數(shù)。使用piercebiotechnologyinc.tm660nm總蛋白質(zhì)分析試劑盒且使用bsa作為標(biāo)準(zhǔn)物來測定反應(yīng)混合物中的酶的量。預(yù)期使abppdc的底物特異性變窄會改進(jìn)特定醛和其下游產(chǎn)物的積聚。一般來說，abppdc偏好更龐大2-酮酸，如5-苯基-2-酮基戊酸酯和苯基丙酮酸，如通過關(guān)于那些底物的高催化效率證明(參見例如spaepen,s.；versees,w.；gocke,d.；pohl,m.；steyaert,j.；vanderleyden,j.“參與巴西固氮螺菌的生長素生產(chǎn)的苯丙酮酸脫羧酶的表征(characterizationofphenylpyruvatedecarboxylase,involvedinauxinproductionofazospirillumbrasilense),”《細(xì)菌學(xué)雜志(j.bacteriol.)》,2007,189:7626-7633)。對表1中列出的abppdc和變異體篩選針對作為底物的2mm2-酮基己酸酯(2-kh)和2mm2-酮基壬酸酯(2-kn)的活性。篩選展現(xiàn)野生型abppdc催化2-kn的脫羧，但在分析條件下展現(xiàn)針對2-kh的不佳活性。所有abppdc變異體也催化2-kn的脫羧，且展現(xiàn)針對2-kh的相對低活性(表1)。gln-536經(jīng)丙氨酸、纈氨酸、異亮氨酸或亮氨酸的取代相比于野生型酶增加2-kn脫羧基活性，但也改進(jìn)針對作為底物的2-kh的活性。這些結(jié)果表明表1中列出的所有abppdc變異體可以活性形式表達(dá)于異源系統(tǒng)中。此外，其全部具有相比于2-kh顯著較高的針對2-kn的活性，表明本文所述的abppdc和變異體偏好>c62-酮酸。對所有酶進(jìn)行詳細(xì)穩(wěn)態(tài)動力學(xué)分析以測定對2-酮基辛酸酯(2-ko)和2-酮基壬酸酯(2-kn)脫羧的最大速率和催化效率。兩種底物均展現(xiàn)如從表2顯而易見的雙曲線和非雙曲線動力學(xué)。對于顯示非雙曲線動力學(xué)的abppdc變異體，將2-ko和2-kn的脫羧的速度擬合至hill方程式(表2圖例)且如表2中所示地計(jì)算最大速率和催化效率(kcat/k0.5)。大于1的希爾系數(shù)表明存在通過2-ko和2-kn的底物活化。已在abppdc和其它脫羧酶的情況下報(bào)導(dǎo)底物活化。另外參見spaepen,s.,同上。如從表2顯而易見，氨基酸取代以不同方式影響捕獲2-ko和2-kn用于催化中的變異體的催化效率。對于一些變異體，例如f532v，2-kn和2-ko的脫羧的催化效率分別為相比于野生型abppdc的180％和45％。這表明f532v取代增加2-kn的底物特異性，同時(shí)減少2-ko的底物特異性。通過采用變異體的催化效率的比率計(jì)算相比于2-ko的abppdc變異體對于2-kn的偏好且顯示于表2中。如表2中所證明，abppdc和f532v的特異性分別為1.8和5.6，指示其對2-kn脫羧的催化效率高于對2-ko脫羧的效率1.8倍和5.6倍。這也指示相比于abppdc，f532v變異體在相比于2-ko偏好2-k中的特異性高3倍。類似地，相比于2-ko的f385l對于2-kn的偏好比abppdc高5倍。此數(shù)據(jù)表明相比于較短2-酮酸(例如2-ko)，f385l和f532v取代改進(jìn)較長2-酮酸(例如2-kn)的底物特異性。因此，當(dāng)使用“+1路徑”延長2-酮酸時(shí)，f385l和f532v變異體將改進(jìn)基于較長(c7-c10)醛的產(chǎn)物的積聚(圖1)。類似地，m461l、f532l、q536g、q536l、f532v/q536v、m380v/m461v、f532a/q536v、f532v/q536a、f385l/q536v、m461v/f532v和m461v/f532v/q536v變異體對于2-kn的特異性相比于每一變異體對于2-ko的特異性分別為3.3、4.3、4.8、2.7、3.6、2.7、6.8、4.6、4.3、5.4和2.1。這表明所有這些變異體在捕獲2-kn用于催化中比abppdc更具特異性。除abppdc變異體對于2-kn的特異性以外，辛醛和衍生自其的生物化學(xué)品的最大積聚也將取決于相對于abppdc變異體的2-kn產(chǎn)生路徑的相對效率。舉例來說，當(dāng)產(chǎn)生2-kn中的工程改造的2-酮酸鏈延長路徑(涉及三種酶，異丙基蘋果酸合成酶、異丙基蘋果酸異構(gòu)酶和異丙基蘋果酸去氫酶)的效率相比于產(chǎn)生2-ko相對較低時(shí)，將由于2-ko通過abppdc變異體的脫羧以及細(xì)胞內(nèi)部基于辛醛的化學(xué)物質(zhì)的積聚的減少而導(dǎo)致庚醛形成。在這些情形下，如f385l的abppdc變異體將基于其相對高特異性(9.1)，以及其減小的作為2-kn脫羧催化劑的效率(表2)而為優(yōu)選的脫羧酶。結(jié)果也顯示經(jīng)疏水性氨基酸(即，甘氨酸、丙氨酸、纈氨酸、亮氨酸或異亮氨酸)取代gln-536改進(jìn)abppdc的催化效率和如表2中所示的其它特異性增強(qiáng)取代。q536v變異體分別在對2-ko和2-kn脫羧中比野生型酶高效8倍和5.7倍(表2)。類似地，m461v/f532v/q536v變異體分別在對2-ko和2-kn脫羧中比m461v/f532v變異體高效27倍和10倍(表2)。m461v/f532v/q536v變異體為分別在對2-ko和2-kn脫羧中比野生型酶高效約17倍和20倍的酶(表2)。m461v/f532v/q536v變異體的較高催化效率允許2-ko在比野生型酶低17倍的胞內(nèi)含量下的有效脫羧且促進(jìn)細(xì)胞內(nèi)部的庚醛衍生性化學(xué)品，如庚醇(經(jīng)由與醇脫氫酶的共表達(dá))或庚酸鹽(經(jīng)由醛去氫酶的共表達(dá))的積聚。也改進(jìn)脫羧的催化效率的gln-536的其它取代(如經(jīng)甘氨酸、丙氨酸、亮氨酸或異亮氨酸)也將改進(jìn)特異性增強(qiáng)取代的催化效率。這通過q536a取代展現(xiàn)，其在添加到f532v變異體(對于2-ko，kcat/k0.5＝4.8mm-1min-1且對于2-kn，kcat/k0.5＝27mm-1min-1)中時(shí)產(chǎn)生針對2-ko和2-kn的催化效率分別高72％和40％的f532v/q536a變異體(對于2-ko，kcat/k0.5＝8.3mm-1min-1和對于2-kn，kcatk0.5＝38mm-1min-1)?？傮w來說，結(jié)果表明abppdc和其基因修飾變異體的表達(dá)允許c7-c11(且在此實(shí)例中尤其為c7-c9)2-酮酸在體內(nèi)的有效脫羧，且進(jìn)而允許例如衍生自醛的化學(xué)物質(zhì)，如己醛、庚醛和/或辛醛在細(xì)胞內(nèi)部的積聚。此外，單獨(dú)或組合形式的f532、f385、q536、m380、m461、f465的修飾可產(chǎn)生在細(xì)胞內(nèi)部展現(xiàn)例如類似地衍生的化學(xué)物質(zhì)的尤其改進(jìn)的積聚的微生物。表2.abppdc和其變異體的動力學(xué)表征**使用本文中描述的htp偶合分析測定初始速度研究。除了通過§指示的所有酶的初始速度使用graphpadprismtm軟件擬合至希爾方程式(v為給定底物濃度s下的初始速度)。kcat、k0.5、h和kcat/k0.5分別為最大速度、最大速度的一半處的底物濃度、希爾系數(shù)和催化效率。結(jié)果為2-3個(gè)獨(dú)立實(shí)驗(yàn)的平均值±標(biāo)準(zhǔn)誤差。通過取用關(guān)于2-kn與2-ko的變異體的催化效率比(kcat/k0.5)計(jì)算abppdc變異體的特異性。1應(yīng)用的命名規(guī)則為第一個(gè)字母指示已改變的氨基酸殘基。f＝苯丙氨酸[phe]；q＝谷氨酰胺[gln]；m＝甲硫氨酸[met]。數(shù)字指示氨基酸序列中的位置(顯示的相應(yīng)為位置380、385、461、465、532和536)。最后一個(gè)字母指示在所述位置處經(jīng)取代的氨基酸殘基。g＝甘氨酸；a＝丙氨酸；i＝異亮氨酸；v＝纈氨酸；l＝亮氨酸。§這些變異體的初始速度擬合至經(jīng)典michaelis-menton方程式。實(shí)例3用巴西固氮螺菌苯丙酮酸脫羧酶(abppdc)的f385l變異體(seqid.8)體外合成c5-c9醇如下進(jìn)行用f385l變異體體外合成直鏈醇：用含0.5mm二磷酸硫胺素、2.5mmnad+、0.2毫克/毫升(0.2mg/ml)牛血清白蛋白(bsa)、5mm乙酰基輔酶a、0.036mg/ml大腸桿菌異丙基蘋果酸合成酶的h97a/s139g/n167g/p169a/g462d變異體(由marcheschi,r.j.等人.“用于碳鏈延長的合成遞歸“+1”路徑(asyntheticrecursive“+1”pathwayforcarbonchainelongation)”《acs化學(xué)生物學(xué)(acschem.biol.)》7:689-697,2012)、0.16mg/ml異丙基蘋果酸異構(gòu)酶的leuc亞單元(基因庫寄存編號nc000913.3，基因標(biāo)識：945076)和0.21mg/ml異丙基蘋果酸異構(gòu)酶的leud亞單元(基因庫寄存編號nc000913.3，基因標(biāo)識：945642)、0.264mg/ml大腸桿菌異丙基蘋果酸去氫酶(leub；基因庫寄存編號nc_000913.3，基因標(biāo)識：944798)、0.192mg/ml異丙基蘋果酸去氫酶的l96g/v198a變異體(報(bào)導(dǎo)于wo2015089127a1中)、0.025mg/ml釀酒酵母醇脫氫酶(adh6，基因庫寄存編號np_014051.3)和0.0054mg/mlf385l變異體(seqid8)的體外合成緩沖液(50mm2-[4-(2-羥乙基)哌嗪-1-基]乙磺酸，ph7.5，含有30mm氯化鉀(kcl)和5mm氯化鎂(mgcl2))培育0.5mm2-酮基丁酸酯(2-kb)。反應(yīng)通過添加2-酮基丁酸酯至反應(yīng)混合物的其余部分而起始。相同體積的分析級甲苯(用于hplc的chromosolvplustm，≥99％，產(chǎn)品目錄編號650579)覆加在反應(yīng)混合物的頂部上且在30℃下培育溶液。在30℃下培育2.5小時(shí)之后，添加nadph至水層直到1mm的最終濃度。在30℃下培育6小時(shí)之后，添加額外nadph至水層直到2mm的最終濃度。在30℃下再培育反應(yīng)物18小時(shí)，接著通過在-20℃下冷凍30分鐘而停止。去除甲苯層的一部分且使用裝備有火焰電離檢測器(fid)的氣相色譜儀分析。通過在以上反應(yīng)混合物中用0.5mm2-酮基異戊酸酯(2-kiv)或0.5mm3-甲基-2-酮基戊酸酯(3m-2kp)置換2-酮基丁酸酯且如上文所述地進(jìn)行實(shí)驗(yàn)而進(jìn)行f385l變異體的分支鏈醇的體外合成。使用裝備有火焰電離檢測器(fid)、g1513a型號自動注射器和gc自動取樣器控制器的hewlettpackard(hp)6890系列氣相色譜儀對醇進(jìn)行定量。使用agilentj&wdb-ffap毛細(xì)管gc柱(30m×0.320mmid×0.25μm膜厚度；產(chǎn)品目錄編號123-3232，agilenttechnologies,inc.,wilmington,de19808)分離分析物。初始gc烘箱溫度為40℃，其保持1.50分鐘，接著以40℃/分鐘梯度增加至235℃。此梯度給出6.38分鐘的總運(yùn)行時(shí)間。柱流動速率為4.0ml/min，其中氦氣作為載氣。注入體積為1μl。注射器和檢測器的溫度設(shè)定為225℃。產(chǎn)生自這些體外合成反應(yīng)的醇效價(jià)顯示于圖2、3和4中。結(jié)果指示f385l變異體與大腸桿菌異丙基蘋果酸合成酶(leua)、大腸桿菌異丙基蘋果酸異構(gòu)酶(leucd)、野生型和經(jīng)修飾大腸桿菌異丙基蘋果酸去氫酶(leub)以及醇脫氫酶(adh6)的h97a/s139g/n167g/-p169a/g462d變異體組合，在與2-酮基丁酸酯、2-酮基異戊酸酯或3-甲基-2-酮基戊酸酯一起培育后產(chǎn)生延長的c5-c9醇。此外，結(jié)果展示f385l變異體對于較長直鏈醇的特異性，其中1-辛醇表示在與2-酮基丁酸酯一起培育后產(chǎn)生的總醇的大致60％。在分支鏈2-酮酸(2-酮基異戊酸酯(kiv)和3-甲基-2-酮基戊酸酯(3m-2-kp))的情況下，在與2-酮基異戊酸酯一起培育后產(chǎn)生大致等效量的5-甲基-1-己醇和6-甲基-1-庚醇，且在與3-甲基-2-酮基戊酸酯一起培育后產(chǎn)生大致等效量的3-甲基-1-戊醇和5-甲基-1-庚醇。這些結(jié)果展示f385l變異體接受直鏈以及分支鏈2-酮酸作為底物且可產(chǎn)生對應(yīng)直鏈和分支鏈醛，其可隨后轉(zhuǎn)化為其它產(chǎn)物，如醇、羧酸或烷烴。實(shí)例4使用野生型abppdc和其變異體以及“+1路徑”酶在大腸桿菌的工程改造菌株中體內(nèi)產(chǎn)生c5-c8醇大腸桿菌(e.coli)mg1655經(jīng)工程改造以促進(jìn)長鏈直鏈醇產(chǎn)生且使得能夠從t7啟動子進(jìn)行基因表達(dá)。為了改進(jìn)直鏈醇生產(chǎn)，ilvbn和ilvih經(jīng)由λred介導(dǎo)的同源重組不活化，如datsenko,ka,wanner,bl,“大腸桿菌k-12中的染色體基因使用pcr產(chǎn)物的單步不活化(one-stepinactivationofchromosomalgenesinescherichiacolik-12usingpcrproducts),”《美國國家科學(xué)院院刊(proc.natl.acad.sci.u.s.a.)》2000,97(12),6640-6645所描述。ilvbn和ilvih基因參與分支鏈氨基酸產(chǎn)生，因此這些基因的不活化消除分支鏈醇的產(chǎn)生。參與2-酮基丁酸酯生產(chǎn)的ilva基因通過經(jīng)由λred介導(dǎo)的同源重組用強(qiáng)力組成性啟動子和強(qiáng)力核糖體結(jié)合位點(diǎn)置換其天然促進(jìn)劑和核糖體結(jié)合位點(diǎn)而上調(diào)，如datsenko和wanner,同上所描述。為了使得能夠從t7啟動子進(jìn)行基因表達(dá)，使用λde3溶原化試劑盒(emdmillipore目錄編號69734)將de3溶源整合到mg1655中。所得菌株基因型為mg1655(de3)δilvbnδilvihilvaup。經(jīng)由八種蛋白質(zhì)的表達(dá)在經(jīng)工程化大腸桿菌菌株中產(chǎn)生c5-c8醇：(1)大腸桿菌異丙基蘋果酸合成酶(leua)；(2)工程改造異丙基蘋果酸合成酶(由marcheschi等人《acs化學(xué)生物學(xué)(acschem.biol.)》2012,7,689-697)；(3)和(4)大腸桿菌異丙基蘋果酸異構(gòu)酶(leucd)的兩個(gè)次單元；(5)異丙基蘋果酸去氫酶(leub)；(6)大腸桿菌異丙基蘋果酸去氫酶的l96g/v198a變異體(如2014年12月10日提交的同在申請中的國際專利申請序號pct/us14/69438(代理人案號75413-wo-pct)中較詳細(xì)描述，所述專利申請要求2013年12月12日提交的美國臨時(shí)專利申請第61/915,040號(代理人案號75413-us-psp)的權(quán)益，兩者均以全文引用的方式并入本文中)；(7)abppdc或其變異體；和(8)釀酒酵母醇脫氫酶(adh6)?？傆?jì)產(chǎn)生十一個(gè)菌株。產(chǎn)生僅含有野生型abppdc的一個(gè)菌株。作為陰性對照，也產(chǎn)生不具有ppdc的菌株。也產(chǎn)生含有abppdc變異體f532v、f358l、f385v、f532vq536v、m461c、m461v、f385vm461c和f385lm461v的八個(gè)菌株。最后，產(chǎn)生含有野生型雷特氏乳球菌(lactococcuslactis)酮基異戊酸酯脫羧酶(kivd；基因寄存編號aj746364)作為比較，因?yàn)橄惹安僮饕扬@示kivd能夠產(chǎn)生與“+1路徑”酶組合的長鏈醇。參見例如marcheschi等人,同上。允許使用四種相容質(zhì)體同時(shí)表達(dá)八種基因的novagenduetvector系統(tǒng)(emdmillipore目錄編號71146、71341、71340和71147)用于表達(dá)上文所提及的基因。四種duet載體中的每一者用t7啟動子下游的八種基因中的兩種克隆，且四種duet載體轉(zhuǎn)化成經(jīng)工程化大腸桿菌菌株。帶有所有質(zhì)體的重組菌株關(guān)于使用抗生素(25微克/毫升(μg/ml)下的安比西林(ampicillin)、17μg/ml下的氯胺苯醇、25μg/ml下的大觀霉素(spectinomycin)和15μg/ml下的康霉素)而選擇且使用所屬領(lǐng)域的技術(shù)人員已知的方法以聚合酶鏈反應(yīng)(pcr)確認(rèn)。在每一固體和液體培養(yǎng)步驟添加抗生素以確保質(zhì)體的維持。在轉(zhuǎn)型、板選擇和pcr確認(rèn)之后，菌株起初在37℃下生長的盧里亞-貝爾塔尼(lb)瓊脂板上培養(yǎng)。單一瓊脂板群落用于接種250ml搖瓶中的50mllb培養(yǎng)基，其使用設(shè)定于200rpm下的恒溫箱振蕩器在37℃下好氧地培養(yǎng)。在lb搖瓶中培養(yǎng)12-16小時(shí)之后，在1％v/v下接種血清瓶以評估醇生產(chǎn)。使用根據(jù)表1中示出的濃度的去離子水制備血清瓶發(fā)酵培養(yǎng)基。培養(yǎng)基經(jīng)過濾滅菌，且將20ml培養(yǎng)基添加至丁基橡膠塞住的125ml血清瓶中。在培養(yǎng)基添加之前，血清瓶通過使用sterisamscocenturysv-160hprevac滅菌器在125℃下高壓滅菌30分鐘而預(yù)滅菌。表1.用于展示來自經(jīng)重組工程改造以含有“+1路徑”以及巴西固氮螺菌脫羧酶(abppdc)或其變異體的大腸桿菌的醇生產(chǎn)的培養(yǎng)基組成。組分濃度(g/l)mops緩沖液26.2甘油20胰蛋白胨10酵母提取物5泛酸鈣1.19na2hpo40.105(nh4)2so40.661nh4cl1.6在接種之后，在37℃下在200rpm于恒溫箱振蕩器中振蕩的情況下培養(yǎng)血清瓶培養(yǎng)物。在接種之后大致三小時(shí)，使用0.1mm異丙基β-d-1-硫代半乳糖苷(iptg)誘導(dǎo)培養(yǎng)物以確保所有基因的表達(dá)。在誘導(dǎo)之后24小時(shí)收獲發(fā)酵物用于分析。在發(fā)酵結(jié)束時(shí)，血清瓶通過置于冷凍機(jī)中20-30分鐘而緊接著冷卻至4℃。將血清瓶去瓶蓋，且將發(fā)酵培養(yǎng)液快速倒入含有1ml飽和氯化鈉溶液和2ml分析級甲苯(用于hplc的chromosolvplustm，≥99.9％，產(chǎn)品目錄編號650579)的50ml錐形管中。培養(yǎng)液-氯化鈉-甲苯混合物經(jīng)渦旋30秒。接著將甲苯萃取物的300μl等分試樣提交至如實(shí)例3中所述地使用gc/fid進(jìn)行分析。血清瓶的平均醇分布顯示于圖5中。結(jié)果指示與“+1路徑”基因和adh6組合的野生型巴西固氮螺菌(abppdc)的表達(dá)導(dǎo)致用于產(chǎn)生介于戊醇至辛醇范圍內(nèi)的直鏈醇的功能路徑。未在無abppdc的菌株中檢測到c5-c8醇，證實(shí)此基因的存在對長鏈醇產(chǎn)生至關(guān)重要。此外，結(jié)果展示含有野生型abppdc的菌株比含有kivd的菌株累積基本上更多的己醇、庚醇和辛醇。未在kivd菌株中檢測到己醇、庚醇或辛醇產(chǎn)生，但abppdcwt菌株分別產(chǎn)生>2mg/l、>3mg/l和>0.1mg/l的己醇、庚醇和辛醇。醇生產(chǎn)的大致50％為庚醇和辛醇，其為相比于主要導(dǎo)致戊醇和己醇生產(chǎn)的其它脫羧酶的情況下的前述成果的顯著改進(jìn)(marcheschi等人《acs化學(xué)生物學(xué)》2012,7:689-697)。因此，使用abppdc脫羧酶似乎將醇生產(chǎn)轉(zhuǎn)化至較長鏈長度，其結(jié)果與實(shí)例1和2內(nèi)包含的體外數(shù)據(jù)一致。圖5中的額外數(shù)據(jù)表明所有abppdc變異體具有產(chǎn)生c5-c8醇的能力。三種abppdc變異體m461c、m461v和f385l/m461v展示相比于野生型abppdc的c5-c8醇產(chǎn)生的顯著改進(jìn)。abppdc變異體m461c產(chǎn)生>3mg/l己醇和>5mg/l庚醇，表示相比于野生型abppdc的大于40％改進(jìn)。最印象深刻地，abppdc變異體m461v顯示相比于野生型abppdc的戊醇、己醇和庚醇產(chǎn)生的大于2倍改進(jìn)。m461v變異體也顯示相對于野生型abppdc的就辛醇效價(jià)來說的30％改進(jìn)。含有m461v變異體的此菌株產(chǎn)生具有約9mg/l己醇和約8.5mg/l庚醇的最高效價(jià)。變異體f532v、f385l、f532vq536v、m461c、m461v和f385lm461v全部顯示相比于kivd和abppdc野生型基因的1-辛醇效價(jià)的改進(jìn)。最后，abppdc變異體f385l/m461v顯示相比于abppdc野生型的己醇和庚醇產(chǎn)生的約60％改進(jìn)。序列表<110>陶氏環(huán)球技術(shù)有限責(zé)任公司p·c·桑格哈尼c·c·斯托爾斯e·c·史樂s·a·格林沃爾特<120>基因修飾苯丙酮酸脫羧酶、其制備方法和用途<130>77272-wo-pct<140>tobeassigned<141>2015-12-03<150>62/089,912<151>2014-12-10<160>82<170>patentinversion3.5<210>1<211>39<212>dna<213>人工序列<220><223>表達(dá)的abppdc變異體的n端處的his-tag的13氨基酸序列<220><221>cds<222>(1)..(39)<400>1atgggcagcagccatcatcatcatcatcacagcagcggc39metglyserserhishishishishishissersergly1510<210>2<211>13<212>prt<213>人工序列<220><223>合成構(gòu)筑體<400>2metglyserserhishishishishishissersergly1510<210>3<211>1638<212>dna<213>人工序列<220><223>野生型巴西固氮螺菌苯丙酮酸脫羧酶（abppdc）<220><221>cds<222>(1)..(1638)<400>3atgaagctggccgaagccctgctgcgcgcgctgaaggatcgcggcgca48metlysleualaglualaleuleuargalaleulysaspargglyala151015caggccatgttcgggattccgggtgatttcgccctgcccttcttcaag96glnalametpheglyileproglyaspphealaleuprophephelys202530gtggcggaggaaacgcagatcctgccgctccacacgctgagccacgag144valalaglugluthrglnileleuproleuhisthrleuserhisglu354045ccggcggtgggcttcgcggcggacgcggcggcgcgttacagttcgacg192proalavalglyphealaalaaspalaalaalaargtyrserserthr505560ctgggggtggcggcggtcacctacggagcgggcgccttcaacatggtg240leuglyvalalaalavalthrtyrglyalaglyalapheasnmetval65707580aacgcggtggccggcgcctacgccgagaagtcgccggtggtcgtcatc288asnalavalalaglyalatyralaglulysserprovalvalvalile859095tccggcgcgccgggcacgacggagggcaacgccggcctgctgctgcac336serglyalaproglythrthrgluglyasnalaglyleuleuleuhis100105110caccagggccgcacgctggacacgcagttccaggtgttcaaggagatc384hisglnglyargthrleuaspthrglnpheglnvalphelysgluile115120125accgtcgcccaggcgcggctggacgacccggccaaggccccggcggag432thrvalalaglnalaargleuaspaspproalalysalaproalaglu130135140atcgcccgcgtgctgggggccgcccgcgccctgtcgcgcccggtctat480ilealaargvalleuglyalaalaargalaleuserargprovaltyr145150155160ctggaaattccccgcaacatggtcaacgccgaggtcgagccggtgggc528leugluileproargasnmetvalasnalagluvalgluprovalgly165170175gacgacccggcttggccggtggaccgcgacgcgctggccgcctgcgcg576aspaspproalatrpprovalaspargaspalaleualaalacysala180185190gacgaggtgctggcggccatgcgctcggccacgtcgccggtgctgatg624aspgluvalleualaalametargseralathrserprovalleumet195200205gtctgcgtcgaggtccgccgctacgggctggaggccaaggtggcggag672valcysvalgluvalargargtyrglyleuglualalysvalalaglu210215220ctggcgcagcggctgggcgtgccggtggtgaccaccttcatggggcgc720leualaglnargleuglyvalprovalvalthrthrphemetglyarg225230235240ggcctgctggccgacgcgccgaccccgccgctcggcacctacatcggc768glyleuleualaaspalaprothrproproleuglythrtyrilegly245250255gtcgccggcgacgcggagatcacccggctggtcgaggagtcggacggg816valalaglyaspalagluilethrargleuvalglugluseraspgly260265270ctgttcctgctcggcgcgatcctcagcgacaccaacttcgcggtgtcc864leupheleuleuglyalaileleuseraspthrasnphealavalser275280285cagcgcaagatcgacctgcgcaagaccatccacgccttcgaccgggcg912glnarglysileaspleuarglysthrilehisalapheaspargala290295300gtgacgctgggctatcacacctacgccgacatcccgctggccgggctg960valthrleuglytyrhisthrtyralaaspileproleualaglyleu305310315320gtggacgccctcttggaagggctgccgccgtccgaccggacgacccgc1008valaspalaleuleugluglyleuproproseraspargthrthrarg325330335ggcaaggagccccacgcctacccgaccggccttcaggcggacggcgag1056glylysgluprohisalatyrprothrglyleuglnalaaspglyglu340345350ccgatcgccccgatggacatcgcccgcgccgtcaacgaccgcgtccgc1104proilealaprometaspilealaargalavalasnaspargvalarg355360365gccgggcaggaaccgctgctgatcgcggcggacatgggcgactgcctg1152alaglyglngluproleuleuilealaalaaspmetglyaspcysleu370375380ttcaccgcgatggacatgatcgacgccgggctgatggcgccgggctat1200phethralametaspmetileaspalaglyleumetalaproglytyr385390395400tacgcgggcatgggcttcggggtgccggcgggcatcggggcgcagtgc1248tyralaglymetglypheglyvalproalaglyileglyalaglncys405410415gtgtcgggcggcaagcgcatcctgaccgtggtcggcgacggcgccttc1296valserglyglylysargileleuthrvalvalglyaspglyalaphe420425430cagatgaccgggtgggagcttggcaactgccgacggctgggcatcgac1344glnmetthrglytrpgluleuglyasncysargargleuglyileasp435440445cccatcgtgatcctgttcaacaacgccagttgggagatgctgcgcacc1392proilevalileleupheasnasnalasertrpglumetleuargthr450455460ttccagcccgaatcggccttcaacgacctggacgactggcgcttcgcc1440pheglnprogluseralapheasnaspleuaspasptrpargpheala465470475480gacatggcggcgggcatgggcggcgacggcgtccgcgtgcgcacacgg1488aspmetalaalaglymetglyglyaspglyvalargvalargthrarg485490495gcggagctgaaggcggcgctggacaaggccttcgccacgcgcgggcgc1536alagluleulysalaalaleuasplysalaphealathrargglyarg500505510ttccagctgatcgaggcgatgatcccgcgcggcgtgctgtccgacacg1584pheglnleuileglualametileproargglyvalleuseraspthr515520525ctggcccgcttcgtccaggggcagaagcgcctgcacgccgcgccccgg1632leualaargphevalglnglyglnlysargleuhisalaalaproarg530535540gagtaa1638glu545<210>4<211>545<212>prt<213>人工序列<220><223>合成構(gòu)筑體<400>4metlysleualaglualaleuleuargalaleulysaspargglyala151015glnalametpheglyileproglyaspphealaleuprophephelys202530valalaglugluthrglnileleuproleuhisthrleuserhis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