本發(fā)明涉及產(chǎn)生3-羥基丙酸(3-hp)的重組酵母和使用其產(chǎn)生3-hp的方法，更具體地，涉及產(chǎn)生3-hp的重組酵母，其包含編碼aadh的外源基因、編碼acc的內(nèi)源基因或外源基因、編碼mcr的外源基因和編碼hpdh的外源基因，并通過[丙酮酸(pyruvate)→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]的生物合成途徑產(chǎn)生3-hp，以及使用該重組酵母產(chǎn)生3-hp的方法。
背景技術(shù)：
：：3-hp(3-羥基丙酸，c3)是乳酸(2-羥基丙酸)的異構(gòu)體，在其兩端具有羧酸基和羥基，因此是能夠轉(zhuǎn)化成各種化學(xué)品(如1,3-丙二醇、丙烯酸、丙烯酰胺、聚合物等)的有用材料。實(shí)際上，由于上述原因，2004年3-hp被美國能源部選為能夠由生物質(zhì)生產(chǎn)的有前途的化學(xué)品之一。特別地，作為用于涂料、粘合劑、添加劑、尿布等的高度市場化的材料，丙烯酸是3-hp的主要應(yīng)用形式。3-hp理論上可以通過發(fā)酵從諸如葡萄糖的各種生物質(zhì)中以100％的收率生產(chǎn)，使用微生物的發(fā)酵工藝適合于滿足對生態(tài)環(huán)境友好和可再生材料的需求。還沒有利用生物質(zhì)商業(yè)生產(chǎn)3-hp的情況，但已經(jīng)對各種方法進(jìn)行了研究，獲得經(jīng)濟(jì)的3-hp產(chǎn)生菌株成為主要障礙。細(xì)菌被公認(rèn)為是產(chǎn)生有機(jī)酸的代表性微生物，并廣泛用于工業(yè)如食品工業(yè)等。然而，將使用工業(yè)細(xì)菌(如大腸桿菌)的有機(jī)酸生產(chǎn)應(yīng)用于大規(guī)?；瘜W(xué)工業(yè)(如3-hp生產(chǎn))存在缺點(diǎn)。隨著有機(jī)酸的產(chǎn)量增加，氫化形式的酸增加，酸性增加(ph降低)，從而大多數(shù)大腸桿菌的活性降低。在生產(chǎn)高濃度的有機(jī)酸的情況下，細(xì)菌需要堿如氫氧化鈉(naoh)和氫氧化銨(nh4oh)來保持中性ph。這導(dǎo)致依賴于注入的堿的發(fā)酵過程的成本增加，使提取和分離過程難以進(jìn)行或者使成本顯著增加。在最近的將有機(jī)酸生產(chǎn)應(yīng)用于化學(xué)工業(yè)的情況中，使用酵母從葡萄糖生產(chǎn)有機(jī)酸。與細(xì)菌相比，酵母對有機(jī)酸具有高耐受性，因此即使在高酸性(低ph)下酵母的活性也不會被顯著抑制，使得酵母成為更合適的用于生產(chǎn)有機(jī)酸的宿主。特別地，酵母已經(jīng)被常規(guī)地、廣泛地用作用于生產(chǎn)烈酒、工業(yè)乙醇等的工業(yè)生物催化劑，酵母可以被大規(guī)模地培養(yǎng)并且可以不被噬菌體污染，使得酵母與細(xì)菌相比在化學(xué)工業(yè)中的適用性更加優(yōu)異。酵母具有產(chǎn)生有機(jī)酸的優(yōu)點(diǎn)，但是使用酵母產(chǎn)生有機(jī)酸如3-hp存在幾個缺點(diǎn)。首先，與細(xì)菌相比，對酵母進(jìn)行基因修飾更為困難，為了表達(dá)特定的代謝酶，應(yīng)當(dāng)鑒定用于表達(dá)代謝途徑以及特定代謝酶的亞細(xì)胞位置，這是由于其細(xì)胞形狀與細(xì)菌不同，被分為細(xì)胞內(nèi)結(jié)構(gòu)體，例如線粒體、過氧化物酶體等。例如，代表性代謝中間體如乙酰輔酶a主要在酵母的線粒體中產(chǎn)生。然而，如果在胞質(zhì)溶膠中產(chǎn)生目標(biāo)產(chǎn)物，則也需要在胞質(zhì)溶膠中產(chǎn)生乙酰輔酶a的方法。此外，由于在真菌界存在大量不同的酵母，并且所有酵母都不滿足對高生產(chǎn)率、耐有機(jī)酸性和大規(guī)模培養(yǎng)的要求，因此應(yīng)當(dāng)有效地選擇適于產(chǎn)生有機(jī)酸的宿主。已知一些酵母有效地從作為己糖的葡萄糖產(chǎn)生乙醇，一些酵母種類也產(chǎn)生有機(jī)酸。在進(jìn)行修飾以使用微生物產(chǎn)生目標(biāo)產(chǎn)物時，重要的是保持代謝反應(yīng)的氧化和還原的整體平衡，并且由葡萄糖發(fā)酵生產(chǎn)乙醇的代謝反應(yīng)是得到適當(dāng)維持的反應(yīng)。即使在基因修飾酵母以產(chǎn)生有機(jī)酸的情況下，也應(yīng)當(dāng)適當(dāng)?shù)乇３稚鲜銎胶?，并且在通過修飾酵母來生產(chǎn)乳酸的情況下，平衡地引入另一還原反應(yīng)的乳酸脫氫酶(ldh)用于補(bǔ)充從乙醛生產(chǎn)乙醇的還原反應(yīng)是重要的。已知在少數(shù)微生物例如橙色綠屈撓菌(chloroflexusaurantiacus)中產(chǎn)生少量3-hp，并且3-hp部分地由微生物如糞產(chǎn)堿桿菌(alcaligenesfaecalis)中的二甲基磺基丙酸分解過程或酵母中尿嘧啶的分解過程形成。通過發(fā)現(xiàn)如上所述的自然界中存在的3-hp，已經(jīng)進(jìn)行了3-hp代謝途徑和相應(yīng)的酶的研究，并且基于最近的研究，進(jìn)行了對以下技術(shù)的研究：通過在大腸桿菌中引入3-hp生物合成所需的基因來產(chǎn)生3-hp或pha的技術(shù)，pha為3-hp的聚合物形式。另外，為了使在自然界中僅作為代謝中間體存在或僅少量產(chǎn)生的3-hp的生產(chǎn)率和產(chǎn)量最大化，必須使用諸如代謝工程技術(shù)、系統(tǒng)生物學(xué)技術(shù)、合成生物學(xué)技術(shù)等的技術(shù)。根據(jù)代謝工程技術(shù)的發(fā)展，預(yù)測使用微生物生產(chǎn)各種物質(zhì)的生產(chǎn)途徑成為可能，從葡萄糖生產(chǎn)3-hp的途徑按照代謝中間體可以大致分為丙烯酰輔酶a途徑、β-丙氨酸途徑、丙二酰輔酶a途徑和甘油途徑。丙烯酰輔酶a途徑是指將從葡萄糖糖酵解獲得的丙酮酸或磷酸烯醇丙酮酸(pep)經(jīng)由乳酸(lactate)或β-丙氨酸轉(zhuǎn)化為丙烯酰輔酶a，然后通過水合反應(yīng)和還原反應(yīng)將丙烯酰輔酶a轉(zhuǎn)化為3-hp的代謝途徑(丙酮酸或pep→乳酸或β-丙氨酸→丙烯酰輔酶a→3-hp)。丙烯酰輔酶a是在丙酸的分解過程中觀察到的代謝物，并且由于形成該代謝物的吉布斯自由能值為正，所以正向反應(yīng)是不利的反應(yīng)。此外，丙烯酰輔酶a硫酯酶的底物特異性低，使得丙烯酰輔酶a途徑不適合作為大規(guī)模生產(chǎn)3-hp的代謝途徑。β-丙氨酸途徑是指通過轉(zhuǎn)氨基反應(yīng)將丙酮酸或草酰乙酸轉(zhuǎn)化為氨基酸，最后通過轉(zhuǎn)氨基反應(yīng)經(jīng)由β-丙氨酸轉(zhuǎn)化為3-hp的代謝途徑(丙酮酸或草酰乙酸→氨基酸→β-丙氨酸→3-hp；us2012/0135481a1)。由于β-丙氨酸向3-hp轉(zhuǎn)化的轉(zhuǎn)氨基反應(yīng)經(jīng)由對微生物具有高毒性的丙二酸半醛進(jìn)行，因此需要具有高活性的3-hp脫氫酶。另外，通常，由于轉(zhuǎn)氨基反應(yīng)形成穩(wěn)態(tài)的氨基酸分子結(jié)構(gòu)的自由基形式，所以該反應(yīng)的酶具有用于減輕自由基的反應(yīng)性的結(jié)構(gòu)。由于該自由基對氧具有強(qiáng)的反應(yīng)性，因此對于平穩(wěn)的轉(zhuǎn)氨基反應(yīng)，基本上需要厭氧條件或用于穩(wěn)定自由基分子的輔酶。丙二酰輔酶a途徑是通過羧化將乙酰輔酶a轉(zhuǎn)化為丙二酰輔酶a，然后通過還原反應(yīng)將丙二酰輔酶a轉(zhuǎn)化為3-hp的代謝途徑(乙酰輔酶a→丙二酰輔酶a→3-hp)，甘油途徑是將葡萄糖轉(zhuǎn)化為甘油，通過脫水反應(yīng)將甘油轉(zhuǎn)化為3-羥基丙醛，然后將3-羥基丙醛轉(zhuǎn)化為3-hp的代謝途徑(葡萄糖→甘油→3-羥基丙醛→3-hp)。由于丙二酰輔酶a途徑和甘油途徑經(jīng)由通常由諸如大腸桿菌等的微生物產(chǎn)生的中間體進(jìn)行，因此主要將這些途徑作為3-hp生產(chǎn)途徑進(jìn)行研究(us2013/0071893a1)。由于可以通過丙二酸(malonate)還原酶和3-hp脫氫酶將丙二酰輔酶a轉(zhuǎn)化為3-hp，并且可以通過甘油脫水酶和醛脫氫酶將甘油轉(zhuǎn)化為3-hp，使用修飾的大腸桿菌將葡萄糖或甘油轉(zhuǎn)化為3-hp的方法是眾所周知的。與轉(zhuǎn)氨基反應(yīng)類似，甘油的脫水反應(yīng)是伴隨有自由基的反應(yīng)，基本上需要輔酶b12以在氧的存在下進(jìn)行反應(yīng)?？紤]到工業(yè)發(fā)酵，由于輔酶(如輔酶b12)的成本，難以使用其作為培養(yǎng)基的物質(zhì)，并且微生物(例如酵母)不能在細(xì)胞中生物合成或吸收相應(yīng)的物質(zhì)，因此β-丙氨酸途徑或甘油途徑不適合作為使用酵母產(chǎn)生3-hp的代謝途徑。最近，已經(jīng)報道了修飾關(guān)鍵酶以克服該問題的研究。[us7,655,451b2]丙二酰輔酶a由胞質(zhì)溶膠中的乙酰輔酶a合成，從而可以還原成3-hp。在細(xì)菌如大腸桿菌的情況下，乙酰輔酶a由胞質(zhì)溶膠中的丙酮酸形成，因此可用作tca循環(huán)或其它代謝反應(yīng)的底物。然而，如上所述，在具有獨(dú)立的亞細(xì)胞區(qū)室的酵母中，通常在線粒體中合成乙酰輔酶a并將其用作tca循環(huán)的底物，并且胞質(zhì)溶膠中的乙酰輔酶a通過乙酸產(chǎn)生反應(yīng)或檸檬酸循環(huán)反應(yīng)來產(chǎn)生，乙酸產(chǎn)生反應(yīng)是乙醇產(chǎn)生反應(yīng)的副反應(yīng)。從乙酸或檸檬酸產(chǎn)生乙酰輔酶a的所有反應(yīng)都是消耗atp的反應(yīng)，并且由于與細(xì)菌相比，酵母還消耗能量以在胞質(zhì)溶膠中獲得乙酰輔酶a，所以酵母在能量學(xué)方面是不利的。在酵母中，諸如胞質(zhì)溶膠的還原狀態(tài)、蛋白質(zhì)合成后的折疊、密碼子使用等的環(huán)境與細(xì)菌中的環(huán)境不同，使得在表達(dá)來自細(xì)菌的外源酶時，不顯示其活性或活性可能顯著降低。此外，由于氧或其它金屬離子的存在或不存在可以顯著改變所述活性，即使在具有相同功能的外源酶的情況下，其在酵母中的表達(dá)結(jié)果會根據(jù)酶的來源而不同。實(shí)際上，在木糖代謝中的重要酶即木糖異構(gòu)酶(xi)的情況下，當(dāng)在酵母中表達(dá)來自細(xì)菌的xi時，通常其活性明顯較低，但是證明了通過引入來自厭氧真菌的xi，酵母被成功修飾從而以相對高的活性進(jìn)行木糖代謝。因此，在酵母中引入來自細(xì)菌或古生菌的代謝途徑(如丙二酰輔酶a途徑)的情況下，應(yīng)通過發(fā)揮相同功能的各種來源的基因來獲得具有高活性的基因。存在與基因修飾各種酵母菌株中的釀酒酵母(saccharomycescerevisiae)以產(chǎn)生3-hp的方法相關(guān)的各種文獻(xiàn)和專利，但是在釀酒酵母的情況下，由于3-hp通過[丙酮酸→乙醛→乙酸→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]途徑生產(chǎn)，存在該產(chǎn)生3-hp的代謝途徑復(fù)雜、3-hp的生產(chǎn)率相對低的問題(us2010/0248233a1；y.chen等人，metabolicengineering，22:104-109,2014)。因此，作為解決上述問題的努力的結(jié)果，為了克服酵母在獲得胞質(zhì)溶膠中乙酰輔酶a的過程中消耗atp的缺點(diǎn)，本發(fā)明人想到更短的代謝途徑：[丙酮酸→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]，該代謝途徑不經(jīng)過乙酸中間體而直接將乙醛轉(zhuǎn)化為乙酰輔酶a，并且證實(shí)在使用包含該途徑的重組酵母的情況下，與使用大腸桿菌的情況不同，不僅減少了ph調(diào)節(jié)物質(zhì)的使用(從而減少了鹽的產(chǎn)生)，而且即使在低的ph下也可以以高濃度和高產(chǎn)率從生物質(zhì)產(chǎn)生3-hp，從而完成本發(fā)明。技術(shù)實(shí)現(xiàn)要素：技術(shù)問題本發(fā)明的目的是提供包含[丙酮酸→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]的主動3-hp生物合成途徑的重組酵母，其不經(jīng)由乙酸而將乙醛直接轉(zhuǎn)化為乙酰輔酶a。本發(fā)明的另一個目的是提供使用該重組酵母產(chǎn)生3-hp的方法。問題的解決方案為了實(shí)現(xiàn)上述目的，本發(fā)明提供了一種包含[丙酮酸→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]的主動3-hp生物合成途徑的重組酵母，其中該酵母包含：編碼aadh的外源基因；編碼acc的內(nèi)源基因或外源基因；編碼mcr的外源基因；和編碼hpdh的外源基因。在本發(fā)明中，所述酵母是耐酸的，并且選自例如酵母屬(saccharomyces)、kazachstania屬和假絲酵母(candida)屬。特別感興趣的酵母種包括釀酒酵母(saccharomycescerevisiae)、kazachstaniaexigua、kazachstaniabulderi和小假絲酵母(candidahumilis)，但種類不僅限于此。此外，本發(fā)明提供了制備3-hp的方法，其包括：(a)在包含至少一種碳源的培養(yǎng)基中培養(yǎng)權(quán)利要求1-10中任一項所述的重組酵母，從而產(chǎn)生3-hp；以及(b)從培養(yǎng)物中分離3-hp。附圖說明圖1示出了本發(fā)明的重組酵母從葡萄糖產(chǎn)生3-hp的途徑(修飾的丙二酰輔酶a代謝途徑)和主要的酶。圖2示出了乙酰輔酶a羧化酶的acc1活性的相對排名。圖3示出了對古細(xì)菌mcr變體的相對活性(左圖)和表達(dá)水平(sds-page)(右圖)的確認(rèn)結(jié)果。圖4示出了用于表達(dá)與3-hp途徑相關(guān)的酶的酵母表達(dá)質(zhì)粒psk-084和psk-085。圖5示出了對可以影響3-hp生產(chǎn)水平的培養(yǎng)條件的測試結(jié)果。圖6示出了使用補(bǔ)料分批(片劑摻加(tabletspiking))培養(yǎng)條件時更成熟的3-hp生產(chǎn)菌株的3-hp生產(chǎn)量。具體實(shí)施方式除非另有定義，本文使用的所有技術(shù)和科學(xué)術(shù)語具有與本發(fā)明所屬領(lǐng)域的技術(shù)人員通常理解的相同的含義。通常，本文使用的命名法是本領(lǐng)域公知的并且通常使用的。通常，在酵母中，由于乙酰輔酶a通過[乙醛→乙酸→乙酰輔酶a]途徑制備，并且在酵母的胞質(zhì)溶膠中產(chǎn)生乙酸，因此在生產(chǎn)乙酰輔酶a的過程中atp通過轉(zhuǎn)化為amp而被消耗掉(圖1；y.chen等人,metabolicengineering,22:104-109,2014)。然而，在本發(fā)明中，設(shè)計并應(yīng)用了一種途徑(圖1)，在產(chǎn)生胞質(zhì)溶膠中的乙酰輔酶a的過程中，該途徑通過不經(jīng)由乙酸而由乙醛直接制備乙酰輔酶a來克服和改善酵母消耗atp的缺點(diǎn)。結(jié)果，在使用本發(fā)明的重組酵母的情況下，證明了即使在低ph下，也以高濃度和高產(chǎn)率由葡萄糖產(chǎn)生3-hp。因此，一方面，本發(fā)明涉及包含[丙酮酸→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]的主動3-hp生物合成途徑的重組酵母，其中該酵母包含：編碼aadh的外源基因；編碼acc的內(nèi)源基因或外源基因；編碼mcr的外源基因；和編碼hpdh的外源基因。[丙酮酸→乙醛→乙酰輔酶a→丙二酰輔酶a→丙二酸半醛→3-hp]的3-hp生物合成途徑是從諸如葡萄糖等的碳源產(chǎn)生3-hp的途徑。“丙酮酸→乙醛”是指使用丙酮酸脫羧酶(pdc)由丙酮酸生產(chǎn)乙醛而不產(chǎn)生中間體的途徑；(ii)“乙醛→乙酰輔酶a”是指使用乙?；胰┟摎涿?acetylatingacetaldehydedehydrogenase，aadh)由乙醛生產(chǎn)乙酰輔酶a而不產(chǎn)生諸如乙酸的中間體的途徑；(iii)“乙酰輔酶a→丙二酰輔酶a”是指使用乙酰輔酶a羧化酶(acc)由乙酰輔酶a生產(chǎn)丙二酰輔酶a而不產(chǎn)生中間體的途徑；(iv)“丙二酰輔酶a→3-hp或丙二酰輔酶a→丙二酸半醛→3-hp”是指使用雙功能丙二酰輔酶a還原酶(mcr)由丙二酰輔酶a生產(chǎn)3-hp而不產(chǎn)生中間體的途徑，或通過使用單功能丙二酰輔酶a還原酶生物合成丙二酸半醛從而由丙二酸半醛產(chǎn)生3-hp的途徑(圖1)。在本發(fā)明中，pdc基因不是工程化的，但是可以例如通過擴(kuò)增酵母中存在的pdc基因、通過應(yīng)用本
技術(shù)領(lǐng)域：
：熟知的現(xiàn)有技術(shù)來工程化pdc基因以增加3-hp生產(chǎn)率。在本發(fā)明的一個示例性實(shí)施方案中，通過生物信息學(xué)基因組挖掘途徑酶候選物來提取編碼具有特殊功能的途徑酶的基因。本發(fā)明包括編碼aadh的外源基因。在一個實(shí)施方案中，編碼aadh的基因?yàn)榫幋a下述aadh的核酸：aadh具有與選自以下表1-3中所示的氨基酸序列的aadh氨基酸序列具有至少60％，優(yōu)選至少70％、80％、90％、91％、92％、93％、94％、95％、96％、97％、99％或100％的序列同一性的氨基酸序列，但編碼aadh的基因不限于此，只要其具有由乙醛生物合成乙酰輔酶a的功能即可。表1aadh(adhe型)表2aadh(eute型)表3aadh本發(fā)明包括編碼acc的基因。在一個實(shí)施方案中，編碼acc的基因?yàn)榫幋a下述acc的核酸，該acc具有與選自下表4所示的氨基酸序列的acc氨基酸序列具有至少60％，優(yōu)選至少70％、80％、90％、91％、92％、93％、94％、95％、96％、97％、99％或100％的序列同一性的氨基酸序列，但編碼acc的基因不限于此，只要其具有由乙酰輔酶a生物合成丙二酰輔酶a的功能即可。表4acc(真核多結(jié)構(gòu)域型)本發(fā)明包括編碼mcr的基因。在一個實(shí)施方案中，mcr可以是雙功能mcr，其具有將丙二酰輔酶a轉(zhuǎn)化為丙二酸半醛的功能和將丙二酸半醛轉(zhuǎn)化為3-hp的功能；或者是單功能mcr，其具有將丙二酰輔酶a轉(zhuǎn)化為丙二酸半醛的功能。在本發(fā)明中，編碼所述雙功能mcr的基因?yàn)榫幋a下述mcr的核酸：mcr具有與選自下表5中所示的氨基酸序列的mcr氨基酸序列具有至少60％，優(yōu)選至少70％、80％、90％、91％、92％、93％、94％、95％、96％、97％、99％或100％的序列同一性的氨基酸序列，但編碼所述雙功能的基因不限于此，只要其同時具有將丙二酰輔酶a轉(zhuǎn)化為丙二酸半醛的功能和將丙二酸半醛轉(zhuǎn)化為3-hp的功能即可。表5mcr(雙功能型)在另一個實(shí)施方案中，所述mcr基因可以是具有將丙二酰輔酶a轉(zhuǎn)化為丙二酸半醛的功能的單功能基因，可以進(jìn)一步包含編碼可以將丙二酸半醛轉(zhuǎn)化為3-hp的酶的基因。在本發(fā)明中，所述編碼單功能mcr的基因?yàn)榫幋a下述mcr的核酸：該mcr具有與選自下表6中所示的氨基酸序列的mcr氨基酸序列具有至少60％，優(yōu)選至少70％、80％、90％、91％、92％、93％、94％、95％、96％、97％、99％或100％的序列同一性的氨基酸序列，但編碼所述單功能mcr的基因不限于此，只要其具有將丙二酰輔酶a轉(zhuǎn)化為丙二酸半醛的功能即可。表6mcr在本發(fā)明中，編碼將丙二酸半醛轉(zhuǎn)化為3-hp的酶的基因是編碼下述酶的核酸：該酶具有與選自下表7-10所示的氨基酸序列的hpdh、hibadh、hbdh或bdh氨基酸序列具有至少60％，優(yōu)選至少70％、80％、90％、91％、92％、93％、94％、95％、96％、97％、99％或100％的序列同一性的氨基酸序列，但所述編碼酶的基因不限于此，只要該蛋白編碼基因具有從丙二酸半醛生物合成3-hp的功能即可。下表7中所示的hpdh氨基酸序列、下表8中所示的hibadh氨基酸序列、下表9中所示的hbdh氨基酸序列和下表10中所示的bdh-氨基酸序列為具有從丙二酸半醛生物合成3-hp的功能的蛋白質(zhì)的氨基酸序列。表7hpdh表8hibadh表9hbdh表10bdh在本發(fā)明的一個具體實(shí)施方案中，包含主動3-hp生物合成途徑的重組酵母可以具有表11中所列的基因，例如編碼具有選自seqidno:74-98的氨基酸序列的aadh的外源基因、編碼具有選自seqidno:99-106的氨基酸序列的acc的內(nèi)源基因或外源基因、編碼具有選自seqidno:107-116的氨基酸序列的mcr的外源基因以及編碼具有選自seqidno：117-144的氨基酸序列的hpdh的外源基因。表11在本發(fā)明中，酵母可以是耐酸的，并且所述耐酸酵母可以選自例如酵母屬、kazachstania屬和假絲酵母屬。特別感興趣的酵母種包括釀酒酵母、kazachstaniaexigua、kazachstaniabulderi和小假絲酵母，但不限于此。根據(jù)本發(fā)明，重組酵母可以是耐酸的，為了制備耐酸重組酵母，優(yōu)選使用對有機(jī)酸(特別是3-hp和/或制備3-hp時作為副產(chǎn)物產(chǎn)生有機(jī)酸)具有耐酸性的酵母宿主。耐酸性酵母可以選自釀酒酵母、kazachstaniaexigua、kazachstaniabulderi和小假絲酵母，但不限于此。本說明書中使用的術(shù)語“耐酸性酵母”是指對有機(jī)酸如3-hp等具有耐酸性的酵母，耐酸性可以通過確認(rèn)在含有各種濃度的有機(jī)酸的培養(yǎng)基上的生長來評價。在這種情況下，“耐酸性酵母”可以是當(dāng)在含有高濃度有機(jī)酸的培養(yǎng)基中生長時與一般酵母相比表現(xiàn)出高生長速率和高生物質(zhì)消耗速率的酵母。根據(jù)本發(fā)明，耐酸性酵母可以是在含有超過1m或更多有機(jī)酸(特別是3-hp)的培養(yǎng)基中、在小于有機(jī)酸(特別是3-hp)的pka值的ph下，與在不含有機(jī)酸的培養(yǎng)基中生長的酵母相比，能夠維持至少10％的葡萄糖(或類似物)消耗速率或至少10％的比生長速率的酵母。根據(jù)本發(fā)明，耐酸性酵母可以是在ph2-4下，與在ph7下相比，可以維持至少10％的葡萄糖(或類似物)消耗速率或至少10％的比生長速率的酵母。本發(fā)明的經(jīng)基因修飾的微生物可以通過將基因插入微生物的染色體或?qū)⒔?jīng)修飾的載體引入微生物中來制備。通常使用dna引入效率高且引入的dna的表達(dá)效率高的宿主作為所述經(jīng)修飾的微生物，在本發(fā)明的一個示例性實(shí)施方案中，使用酵母，但不限于此，只要能充分表達(dá)靶dna，可以使用任何種類的微生物。所述經(jīng)修飾的微生物可以通過任何轉(zhuǎn)化方法制備?！稗D(zhuǎn)化”是指將dna引入宿主，從而dna能夠作為染色體的因子或通過整合染色體而復(fù)制，這是人為引起基因改變的現(xiàn)象。在常見的轉(zhuǎn)化方法中，存在電穿孔、乙酸鋰-peg方法等。另外，在本發(fā)明中，可以使用一般公知的基因工程方法作為向宿主微生物的染色體中引入基因的方法，例如存在使用逆轉(zhuǎn)錄病毒載體、腺病毒載體、腺伴隨病毒載體、單純皰疹病毒載體、痘病毒載體、慢病毒載體、非病毒載體等的方法?！拜d體”是指包含待可操作地連接到合適的控制序列的、能夠在宿主內(nèi)部表達(dá)dna的dna序列的dna構(gòu)建體。載體可以是質(zhì)粒、噬菌體顆?；蚝唵蔚臐撛诨蚪M插入片段。當(dāng)載體轉(zhuǎn)化到合適的宿主中時，其可以被復(fù)制或發(fā)揮功能而不管宿主基因組如何，或在一些情況下，其可以整合到基因組本身中。質(zhì)粒是最常用作載體的類型?？捎糜谠撃康牡牡湫唾|(zhì)粒載體的結(jié)構(gòu)包含(a)允許有效進(jìn)行復(fù)制以使每個宿主細(xì)胞包含質(zhì)粒載體的復(fù)制起點(diǎn)，(b)抗生素抗性基因或營養(yǎng)缺陷型標(biāo)記基因，其允許選擇用質(zhì)粒載體轉(zhuǎn)化的宿主細(xì)胞，和(c)可以插入外源dna片段的限制酶限制性位點(diǎn)。即使不存在合適的限制酶限制性位點(diǎn)，當(dāng)使用根據(jù)一般方法的接頭或合成的寡核苷酸銜接子(adaptor)時，可以容易地將載體和外源dna連接。當(dāng)核酸與其他核酸序列以功能關(guān)系排列時，其是“可操作地連接的”。它可以是：當(dāng)適當(dāng)分子(例如轉(zhuǎn)錄激活蛋白)連接到控制序列時在能夠進(jìn)行基因表達(dá)的過程中連接的基因和控制序列。例如，當(dāng)表達(dá)參與多肽分泌的前蛋白時，前序列或分泌前導(dǎo)序列的dna可操作地連接至多肽的dna；在影響序列的轉(zhuǎn)錄時啟動子或增強(qiáng)子可操作地連接至編碼序列；當(dāng)影響序列的轉(zhuǎn)錄時，核糖體結(jié)合結(jié)構(gòu)域可操作地連接至編碼序列；或當(dāng)核糖體結(jié)合結(jié)構(gòu)域被設(shè)置為易于被翻譯時，其與編碼序列可操作地連接。通常，“可操作地連接”是指連接的dna序列的接觸，或分泌前導(dǎo)序列在前導(dǎo)框中接觸和呈遞。然而，增強(qiáng)子不需要接觸。增強(qiáng)子序列的連接通過在方便的限制酶位點(diǎn)處的連接來進(jìn)行。當(dāng)不存在該結(jié)構(gòu)域時，使用根據(jù)一般方法的合成寡核苷酸銜接子或接頭。當(dāng)然，應(yīng)當(dāng)理解，所有載體都不能等同地發(fā)揮作用以表達(dá)根據(jù)本發(fā)明的dna序列。同樣地，所有宿主細(xì)胞對于相同的表達(dá)系統(tǒng)也不是等同地起作用。然而，在不脫離本發(fā)明的范圍且沒有過度的實(shí)驗(yàn)的情況下，本領(lǐng)域技術(shù)人員可以適當(dāng)?shù)剡x擇載體、表達(dá)控制序列和宿主細(xì)胞。例如，在選擇載體時，必須考慮宿主細(xì)胞。這是因?yàn)檩d體應(yīng)當(dāng)在宿主細(xì)胞中復(fù)制。此外，還應(yīng)當(dāng)考慮復(fù)制數(shù)量和控制載體的復(fù)制數(shù)量的能力以及由載體編碼的其它蛋白質(zhì)(例如抗生素標(biāo)記物)的表達(dá)。在另一方面，本發(fā)明涉及制備3-hp的方法，其包括：(a)在包含至少一種碳源的培養(yǎng)基中培養(yǎng)權(quán)利要求1-10中任一項所述的重組酵母，從而產(chǎn)生3-hp；以及(b)從培養(yǎng)物中分離3-hp。在本發(fā)明中，碳源可以是選自葡萄糖、木糖、阿拉伯糖、蔗糖、果糖、纖維素、葡萄糖低聚物和甘油中的一種或多種，但不限于此。在本發(fā)明中，培養(yǎng)優(yōu)選在微生物如大腸桿菌不再工作(例如產(chǎn)生代謝物等)的條件下進(jìn)行。在實(shí)施方案中，在ph1.0-6.5，優(yōu)選在ph1.0-6.0，更優(yōu)選在ph2.6-4.0下進(jìn)行培養(yǎng)，但不限于此。具體實(shí)施方式在下文中，將參考以下實(shí)施例更詳細(xì)地描述本發(fā)明。然而，以舉例的方式提供以下實(shí)施例是為了容易地解釋對本發(fā)明技術(shù)精神的描述和本發(fā)明技術(shù)精神的范圍。因此，本發(fā)明的范圍不限于此或不由此改變。另外，本發(fā)明所屬領(lǐng)域的技術(shù)人員可根據(jù)本說明書的描述進(jìn)行各種修改和改變。實(shí)施例1基于對3-hp的耐受性選擇宿主酵母菌株生產(chǎn)3-hp的生物體的基本特征是對高濃度3-hp的良好耐受性，這使得在發(fā)酵期間產(chǎn)物能夠積累，同時菌株性能損失最小。對一個718個野生型酵母菌株組成的大的、多樣化的組(set)進(jìn)行篩選以鑒定在低ph下能夠耐受高濃度3-hp并且在這些條件下可以生長和代謝葡萄糖的那些菌株(表12)。最初使用許多基于瓊脂板和液體培養(yǎng)基微量滴定板的生長測定來篩選整組菌株的耐酸性。然后，評價菌株亞組(subset)在搖瓶培養(yǎng)中在低ph下耐受大量3-hp的能力。這些篩選方法中沒有一個孤立地是工業(yè)環(huán)境中3-hp耐受性的完美指標(biāo)，但是許多方法的組合提供了一種全面而穩(wěn)健的確定有前途的產(chǎn)生3-hp酵母菌株的方法。最初，在含有不同量的3-hp的固體ypd基瓊脂培養(yǎng)基上評估718個酵母菌株的生長，其中3-hp的量為：0g/l3-hp(ph6.62)、50g/l3-hp(ph3.44)、75g/l3-hp(ph3.28)、100g/l3-hp(ph3.17)和125g/l3-hp(ph3.08)。然后基于它們在該篩選測定中耐受不同量的3-hp的能力對菌株進(jìn)行評分。然后使用可以自動孵育、搖動和測量培養(yǎng)物濁度的bioscreenc機(jī)器在不存在3-hp(基于scd的培養(yǎng)基，0g/l3-hp，ph6.0)或存在3-hp(基于scd的培養(yǎng)基，70g/l3-hp，ph3.5)的情況下在微量滴定板液體培養(yǎng)物中評估718個酵母菌株的生長。然后采用現(xiàn)有的用于建模微生物生長曲線的軟件在每個實(shí)驗(yàn)條件下為每個菌株確定滯后期、最大生長速率和最終細(xì)胞密度。對于該篩選測定，基于其在不存在3-hp時的最大生長速率、其在3-hp存在下的最大生長速率和這兩個最大生長速率值之間的相對差對每個菌株進(jìn)行評分。使用液體處理自動控制儀器(robot)，在微量滴定板中在含有85g/l3-hp(ph3.5)的基于ypd的液體培養(yǎng)基中評估718個酵母菌株的生長和葡萄糖利用率。使用自動化工作流程(work-flow)來接種生長平板、在指定時間點(diǎn)稀釋樣品用于od測量并離心和收集上清用于hplc分析，hplc分析用于測定指定時間點(diǎn)處殘留葡萄糖量。與bioscreenc生長測定相反，自動操作(robotic)微量滴定板生長測定允許實(shí)現(xiàn)更多的通氣和更高的最大細(xì)胞密度，同時還允許評估葡萄糖利用率，而不像之前的兩種測定一樣僅評估生長速率。對于該篩選測定，基于在3-hp存在下獲得的菌株最大細(xì)胞密度和菌株在3-hp存在下在低ph條件下消耗葡萄糖的能力對每個菌株進(jìn)行評分。將來自各種3-hp耐受性評估測定的個體評分一起取平均值以獲得每種菌株的最終3-hp耐受性評分(表12)。這些篩選表明，在低ph和不同條件下，以下酵母種類具有對3-hp的良好的一般耐受性：蜂生假絲酵母(candidaapicola)、小假絲酵母、東方伊薩酵母(issatchenkiaorientalis)、kazachstaniabulderi、kazachstaniaexigua、膜璞畢赤酵母(pichiamembranifaciens)、釀酒酵母和解脂耶氏酵母(yarrowialipolytica)。然后在搖瓶培養(yǎng)中在低ph下進(jìn)一步分析這8種3-hp耐受性酵母種對3-hp的耐受性。對于這些培養(yǎng)(定義的基于scd的培養(yǎng)基，高初始生物質(zhì)，低通風(fēng))，在以下不同的ph值和不同水平的3-hp存在下評估菌株生長、消耗葡萄糖和產(chǎn)生乙醇的能力：100g/l3-hp(ph4.0)、100g/l3-hp(ph3.5)、100g/l3-hp(ph3.0)和80g/l3-hp(ph2.6)。這些搖瓶培養(yǎng)顯示某些小假絲酵母(c.humilis)、k.bulderi、k.exigua和釀酒酵母菌株在低ph下對高水平的3-hp具有非常強(qiáng)的耐受性，因?yàn)樵诟鞣N酵母菌株中它們在這些苛刻條件下具有最快的葡萄糖利用率、生物質(zhì)產(chǎn)生速率和乙醇產(chǎn)生速率。另一方面，在這些非常受限的生長條件下，蜂生假絲酵母、東方伊薩酵母、膜璞畢赤酵母和解脂耶氏酵母菌株不能良好地表現(xiàn)。這些使用各種搖瓶培養(yǎng)的詳細(xì)后續(xù)分析證實(shí)某些小假絲酵母、k.bulderi、k.exigua和釀酒酵母菌株顯示出作為3-hp產(chǎn)生宿主的巨大潛力，因?yàn)樗鼈冿@示出在工業(yè)相關(guān)條件下對3-hp的高的天然耐受性。表12酵母菌株對3-hp的耐受性實(shí)施例2生物信息學(xué)基因組挖掘以獲得途徑酶候選物進(jìn)行基于同源性的數(shù)據(jù)庫搜索以鑒定用于特定功能的候選酶。在每種情況下，用多個查詢序列搜索數(shù)據(jù)庫。此外，相關(guān)蛋白家族的成員基于interpro結(jié)構(gòu)域注釋從uniprot/swissprot中檢索。使用blastp針對uniprot(swissprot和trembl)和genbank蛋白質(zhì)數(shù)據(jù)庫(nr，pat和env_nr)，以及使用tblastn針對genbank核苷酸數(shù)據(jù)庫(tsa_nt，env_nt和pat)進(jìn)行基于同源性的搜索。提取e值小于1e-30的序列；然而，在一些情況下，對e值小于1e-80的序列進(jìn)行額外的分析。使用查詢序列作為翻譯中的指導(dǎo)，用genewise將命中的核苷酸翻譯成蛋白質(zhì)序列。對于genewise而言，翻譯長acc序列的任務(wù)太困難，因此，從blast-xml輸出中提取蛋白質(zhì)序列(與查詢序列匹配的部分)。為了去除冗余序列，使用blastclust或cd-hit將檢索的序列集中(clustered)成含有彼此具有80％以上同一性的序列的集合(cluster)。從每個集合僅保留一個代表性序列。將非冗余的序列組要么與蛋白質(zhì)家族的pfam結(jié)構(gòu)域比對要么通過使用mafft進(jìn)行比對。在蛋白質(zhì)家族不與任何pfam相關(guān)的情況下或者如果蛋白質(zhì)的序列被分割成幾個pfam結(jié)構(gòu)域，則創(chuàng)建mafft的全局比對?；谑褂胮hylip或fasttree的多序列比對生成系統(tǒng)發(fā)生樹。該樹用e.c.數(shù)、生物體名稱、blaste值注釋，并使用geneious軟件可視化。2-1：乙?；胰┟摎涿敢胰┻€原為乙酰輔酶a可以通過乙?；胰┟摎涿?aadh，e.c.1.2.1.10)完成。aadh可以分為三組功能同源物(wei等人，nat.commun.4：2580,2013)，包括1)具有aadh和醇脫氫酶活性的雙功能蛋白(大腸桿菌adhe型基因，genbankno.：np_415757，查詢序列)，2)參與乙醇胺分解代謝的蛋白質(zhì)(大腸桿菌eute型基因，genbankno.：aag57564，查詢序列)和3)雙功能蛋白質(zhì)，其是參與4-羥基-2-酮戊酸分解代謝的醛縮酶-脫氫酶復(fù)合物的一部分(大腸桿菌mphf型基因，genbankno.：np_414885)。特別感興趣的是第1組(adhe型)和第2組(eute型)的酶。adhe蛋白的n-末端結(jié)構(gòu)域與醛:nad+氧化還原酶高度同源，而c末端區(qū)與fe2+-依賴性乙醇:nad+氧化還原酶家族同源(membrillo-hernandez等人，j.biol.chem.275:33869-33875,2000)。乙?；胰┟摎涿富钚砸部梢酝ㄟ^去除醇脫氫酶結(jié)構(gòu)域來截短雙功能adhe蛋白從而僅具有n-末端醛還原酶結(jié)構(gòu)域而引入細(xì)胞。基于生物信息學(xué)分析和序列同源性推斷具有這種雙功能aadh活性的其他基因。各種基因總結(jié)在表1中。許多腸桿菌可以利用乙醇胺作為碳源和氮源(stojiljkovic等人，j.bacteriol.177:1357-1366,1995)。該分解代謝途徑包括通過乙?；胰┟摎涿竐ute將乙醛轉(zhuǎn)化為乙酰輔酶a的步驟?；谏镄畔W(xué)分析和序列同源性推斷具有這種aadh活性的新基因。各種基因總結(jié)在表2中。此外，基于生物信息學(xué)分析和與adhe以及eute型基因的序列同源性，存在一組注釋為醛脫氫酶的基因，可以推斷其具有aadh活性。各種基因總結(jié)在表3中。2-2：真核乙酰輔酶a羧化酶乙酰輔酶a羧化酶(acc，ec6.4.1.2)是多功能生物素依賴性羧化酶，是脂肪酸生物合成的關(guān)鍵酶。它使用輔因子atp和生物素催化乙酰輔酶a轉(zhuǎn)化為丙二酰輔酶a。該反應(yīng)分兩步進(jìn)行。首先，生物素羧化酶催化生物素與碳酸氫鹽的atp依賴性羧化。然后，羧基轉(zhuǎn)移酶將羧基從生物素轉(zhuǎn)移到乙酰輔酶a以形成丙二酰輔酶a。真核生物酶是大的多結(jié)構(gòu)域酶，而相應(yīng)的原核生物酶由由不同基因編碼的多個亞基組成。acc的活性在轉(zhuǎn)錄水平上以及在轉(zhuǎn)錄后水平上受控(例如通過磷酸化和聚集)以維持乙酰輔酶a動態(tài)平衡。在不同酵母種中的acc工程化導(dǎo)致acc活性增加和丙二酰輔酶a衍生產(chǎn)物的產(chǎn)量增加。已證明或假定在釀酒酵母(genbankno.：caa96294.1，查詢序列)、解脂耶氏酵母(genbankno.：xp_501721.1，查詢序列)和卷枝毛霉(mucorcircinelloides)(genbankno.：epb82652.1，查詢序列)中存在編碼具有acc活性的酶的基因。在kazachstaniaexigua(seqidno：1)和小假絲酵母(seqidno：2)的新測序的基因組中鑒定候選乙酰輔酶a羧化酶基因并克隆到我們的酵母表達(dá)質(zhì)粒中?；谏镄畔W(xué)分析和序列同源性推斷具有acc活性的其它基因。各種真核多結(jié)構(gòu)域acc基因總結(jié)在表4中。2-3：雙功能丙二酰輔酶a還原酶丙二酰輔酶a還原成3-hp(經(jīng)由丙二酸半醛中間體)可以通過大的雙功能丙二酰輔酶a還原酶完成，其具有c末端醛脫氫酶結(jié)構(gòu)域和n-末端醇脫氫酶結(jié)構(gòu)域的雙功能。具有這種活性的、高度底物特異性的、nadph依賴性酶的特征在于參與稱為3-羥基丙酸循環(huán)的自養(yǎng)co2固定途徑的光營養(yǎng)型綠色無硫細(xì)菌橙色綠屈撓菌(chloroflexusaurantiacus)(genbankno.：aas20429.1；查詢序列)(hugler等人，j.bacteriol.184:2404-2410,2002)?；谏镄畔W(xué)分析和序列同源性推斷具有該雙功能丙二酰輔酶a還原酶活性的其他基因。各種基因總結(jié)在表5中。2-4：丙二酰輔酶a還原酶與上面討論的雙功能丙二酰輔酶a還原酶相反，丙二酰輔酶a也可以通過兩種單獨(dú)的酶催化成3-hp。通過該途徑，丙二酰輔酶a首先通過丙二酰輔酶a還原酶(mcr；ec1.2.1.75)或輔酶a?；岚肴┟摎涿高€原為丙二酸半醛，然后通過3-羥基丙酸脫氫酶(3-hpdh；ec1.1.1.59或ec1.1.1.298)還原為3-hp。mcr是一種nadph依賴性酶，其被一些嗜熱嗜酸性古細(xì)菌用于經(jīng)由3-羥基丙酸/4-羥基丁酸循環(huán)自養(yǎng)地將碳固定到有機(jī)物質(zhì)中(berg等人，science，318:1782-1786,2007)。編碼具有這種mcr活性的酶的基因被表征存在于勤奮金屬球菌(metallosphaerasedula)(genbankno.：abp94884.1，查詢序列)和頭寇岱硫化葉菌(genbankno.：bab67276.1，查詢序列)中。雖然這些mcr都具有與橙色綠屈撓菌雙功能丙二酰輔酶a還原酶類似的醛脫氫酶活性，但它們沒有表現(xiàn)出任何顯著的序列相似性，表明綠屈撓菌屬和硫化葉菌科中的自養(yǎng)途徑趨同進(jìn)化，并且募集不同的基因以在這些分類群中執(zhí)行相似的代謝過程(alber等人，j.bacteriol.188:8551-8559,2006)。特別地，古細(xì)菌mcr顯示出與天冬氨酸-半醛脫氫酶的高的序列相似性?；谏镄畔W(xué)分析和序列同源性推斷具有該mcr活性的其他基因。各種基因總結(jié)在表6中。2-5：3-羥基丙酸脫氫酶丙二酸半醛可通過可逆的3-羥基丙酸脫氫酶(hpdh；ec1.1.1.59，nadh依賴性的)或丙二酸半醛還原酶(ec1.1.1.298，nadph依賴性的)還原為3-hp。這些酶天然參與細(xì)菌和植物中的β-丙氨酸代謝、丙酸代謝或尿嘧啶降解。此外，一些嗜熱嗜酸性古細(xì)菌需要這些酶以通過3-羥基丙酸/4-羥基丁酸循環(huán)固定碳(kockelkorn和fuchs，j.bacteriol.191:6352-6362,2009)。已證明或假定在大腸桿菌(genbankno.：efv00080.1，查詢序列)、釀酒酵母(genbankno.：daa10125.1，查詢序列)、勤奮金屬球菌(genbankno.：abp96133.1，查詢序列)、頭寇岱硫化葉菌(genbankno.：bak54608.1，查詢序列)和大腸桿菌(genbankno.：acr64730.1，查詢序列)中存在編碼具有3-羥基丙酸脫氫酶或丙二酸半醛還原酶活性的酶的基因。基于生物信息學(xué)分析和序列同源性推斷具有3-羥基丙酸脫氫酶或丙二酸半醛還原酶活性的其它基因。各種基因總結(jié)在表7中。2-6：3-羥基異丁酸脫氫酶3-羥基異丁酸脫氫酶(hibadh；ec1.1.1.31)是參與纈氨酸和其它支鏈氨基酸的代謝的關(guān)鍵酶。hibadh催化3-羥基異丁酸向甲基丙二酸半醛的nadh依賴性或nadph依賴性可逆轉(zhuǎn)化。然而，由于其廣泛的底物特異性，還已表明hibadh通過將丙二酸半醛轉(zhuǎn)化為3-hp顯示出3-羥基丙酸脫氫酶活性(即ec1.1.1.59)(yao等人，appl.bio.biotechnol.160:694-703,2010)。在惡臭假單胞菌(genbankno.：adr61938.1，查詢序列)、綠膿桿菌(genbankno.：aag06957.1，查詢序列)、蠟樣芽胞桿菌(genbankno.：aap10961.1，查詢序列)和糞產(chǎn)堿桿菌(genbankno.：ejc65559.1，查詢序列)中鑒定了具有hibadh活性的酶。基于生物信息學(xué)分析和序列同源性推斷具有該hibadh活性的其他基因。各種基因總結(jié)在表8中。2-7：4-羥基丁酸脫氫酶4-羥基丁酸脫氫酶(hbdh；ec1.1.1.61)是天然參與丁酸代謝的酶。hbdh催化4-羥基丁酸向琥珀酸半醛的可逆nad+依賴性轉(zhuǎn)化。然而，因?yàn)槊阜磻?yīng)相似，hbdh也可以將丙二酸半醛轉(zhuǎn)化為3-hp。已經(jīng)在鉤蟲貪銅菌(genbankno.：aac41425.1，查詢序列)和克氏梭菌(genbankno.：edk35022.1，查詢序列)中鑒定了具有hbdh活性的酶?；谏镄畔W(xué)分析和序列同源性推斷具有該hbdh活性的其它基因。各種基因總結(jié)在表9中。2-8：3-羥基丁酸脫氫酶3-羥基丁酸脫氫酶(bdh；ec1.1.1.30)是天然參與丁酸代謝的酶。bdh催化3-羥基丁酸向乙酰乙酸酯的可逆nad+依賴性轉(zhuǎn)化，但其也可氧化其它3-羥基一元羧酸。例如，因?yàn)槊阜磻?yīng)是類似的，bdh可以將丙二酸半醛轉(zhuǎn)化為3-hp。已經(jīng)在綠膿桿菌(genbankno.：gaa17557.1，查詢序列)中鑒定了具有bdh活性的酶?；谏镄畔W(xué)分析和序列同源性推斷具有該bdh活性的其他基因。各種基因總結(jié)在表10中。實(shí)施例3酶活性的測量acc分光光度酶測定分光光度acc測定是偶聯(lián)測定，其中由acc反應(yīng)產(chǎn)生的產(chǎn)物在需要輔因子nad(p)h的反應(yīng)中被進(jìn)一步消耗掉，可以用分光光度計監(jiān)測輔因子nad(p)h的氧化。kroeger等人(2011，anal.biochem.411：100-105)描述了偶聯(lián)測定，其中由acc1產(chǎn)生的丙二酰輔酶a通過純化的丙二酰輔酶a還原酶(mcr)在需要nadph作為輔因子的反應(yīng)中進(jìn)一步轉(zhuǎn)化為丙二酸半醛。acc活性通過以下nadph氧化來測量。diacovich等人(2002，j.biol.chem.277:31228-31236)將adp轉(zhuǎn)化(在acc反應(yīng)中用作輔因子的atp的水解產(chǎn)物)與需要adp的丙酮酸激酶反應(yīng)結(jié)合，該丙酮酸激酶反應(yīng)進(jìn)一步與使用乳酸脫氫酶形成丙酮酸的反應(yīng)偶聯(lián)。后一種酶需要nadh作為輔因子，隨后該輔因子被氧化。acc放射性酶測定最常用的體外acc測定是基于放射性14c碳酸鹽的使用。隨后是將放射性碳酸鹽摻入酸和非揮發(fā)性物質(zhì)(即丙二酰輔酶a)中。通過酸和熱處理將未轉(zhuǎn)化為丙二酰輔酶a的14c標(biāo)記的碳酸氫鈉除去，酸和熱處理將剩余的nah14co3和可能的反應(yīng)副產(chǎn)物轉(zhuǎn)化為14c標(biāo)記的co2。經(jīng)過略微的改變，已將由diacovich等人(2002,j.biol.chem.277:31228-31236)描述的測定用于檢測酵母溶解產(chǎn)物的acc活性(shi等人2014,mbio5:3e01130-14)。由在指數(shù)期晚期或穩(wěn)定期收獲的酵母細(xì)胞制備細(xì)胞溶解產(chǎn)物。洗滌細(xì)胞，然后重懸于含有100mmph7.5的磷酸鉀、2mmmgcl2、1mm二硫蘇糖醇和1×edta游離的完整蛋白酶抑制劑(roche)的裂解緩沖液中。通過玻璃珠破碎細(xì)胞，并在4℃下離心后收集上清液。acc酶測定反應(yīng)混合物包括100mm磷酸鉀(ph8.0)、300μgbsa、3mmatp、5mmmgcl2、10mmnah4co3[比活性200μcimmol-1(7400kbqmmol)]和0.5mm乙酰輔酶a。該反應(yīng)的總體積為100μl，包括20μl的細(xì)胞提取物。將反應(yīng)在30℃下孵育15分鐘，并通過加入50μl的5mhcl終止反應(yīng)。將管內(nèi)容物在95℃下蒸發(fā)至干，使殘余物重懸于100μl水中，并與3ml閃爍混合液(scintillationcocktail)(ultimagoldab，perkinelmer)混合。使用液體閃爍計數(shù)器(perkinelmertri-carb2810tr)測定樣品的14c含量。aadh酶測定按照kozak等人(2014,metab.eng.21:46-59)的描述通過在30℃下在340nm處監(jiān)測nad+的還原來測量aadh活性。收集用于獲得細(xì)胞溶解產(chǎn)物的酵母細(xì)胞，用水洗滌，然后重懸于含有100mmtris-hcl緩沖液(ph7.5)和1×edta游離蛋白酶抑制劑混合物(roche)的裂解緩沖液中。將細(xì)胞在precellys24均質(zhì)器中以5500rpm的速度用玻璃珠破碎3×40秒，在各輪破碎之間將細(xì)胞保持在冰上。將溶解產(chǎn)物于4℃下以16000g離心20分鐘，收集上清液。使用bradford方法測定總蛋白濃度。酶測定反應(yīng)混合物在200μl的總反應(yīng)體積中含有0.1mm輔酶a、50mmches緩沖液(ph9.5)、0.8mmnad+、0.2mmdtt和10μl細(xì)胞提取物。通過加入10mm新制備的乙醛溶液開始反應(yīng)，隨后使用thermokonelab20xt分析儀跟蹤nad+的還原。酵母細(xì)胞溶解產(chǎn)物的mcr酶測定根據(jù)經(jīng)過略微改變的chen等人(2014，metab.eng.22:104-109)描述的方法測量mcr酶活性。該方法以在340nm處監(jiān)測nad(p)h的氧化為基礎(chǔ)。收集細(xì)胞并用含有20mmtris-hcl(ph7.5)、20mmnan3的冷的洗滌緩沖液洗滌，然后重懸浮于含有50mmhepes(ph7.5)、150mmkcl、1mmdtt、1mmedta、0.5％tritonx-100和1×edta游離蛋白酶抑制劑混合物(roche)的1ml破碎緩沖液。將細(xì)胞在precellys24均質(zhì)器中以5500rpm的速度用玻璃珠破碎3×40秒，在各輪破碎之間將細(xì)胞保持在冰上。將溶解產(chǎn)物在4℃下以16000g離心20分鐘，收集上清液。使用bradford方法測定總蛋白濃度。mcr測定混合物含有50mmtris-hcl緩沖液(ph8.0)、5mmmgcl2和0.3mmnadph或nadh。在將20μl細(xì)胞溶解產(chǎn)物加入到200μl的總反應(yīng)體積中后，將反應(yīng)在30℃下預(yù)孵育5分鐘，之后通過加入0.15mm丙二酰輔酶a開始反應(yīng)。使用thermokonelab20xt分析儀在340nm監(jiān)測測定。酵母表達(dá)載體的產(chǎn)生產(chǎn)生一系列酵母表達(dá)質(zhì)粒以評估候選基因在酵母中的表達(dá)和活性能力。首先，設(shè)計了新的基于pbluescript的多克隆位點(diǎn)(mcs)，使得可以利用所有可能的限制酶(re)位點(diǎn)組合。然后將該修飾的mcs置于基于prs的系列酵母著絲粒和多拷貝質(zhì)粒中。之后，使用一組10個獨(dú)特的re位點(diǎn)，將九個不同組的啟動子和終止子克隆到這些基于prs的酵母表達(dá)載體中。因此，可以利用該質(zhì)粒系統(tǒng)在合適的酵母菌株中要么以低拷貝數(shù)要么以高拷貝數(shù)同時組成型表達(dá)多達(dá)9個基因，以評價用于產(chǎn)生3-hp的多種途徑酶組合。acc基因的克隆已經(jīng)轉(zhuǎn)化到具有和不具有snf1缺失的工業(yè)釀酒酵母vsk-128菌株的acc1基因示于表13中。acc基因從多拷貝質(zhì)粒表達(dá)，其中它們在pdc1啟動子的控制下。在shi等人的出版物(2014)中描述的突變acc1sc基因是用quikchangeii定點(diǎn)誘變試劑盒(安捷倫科技有限公司)構(gòu)建的。表13轉(zhuǎn)化到釀酒酵母vsk-128的乙酰輔酶a羧化酶基因acc酶測定在文獻(xiàn)中，分光光度測定法已經(jīng)用于用純化的或部分純化的acc酶檢測acc酶活性。用分光光度測定測試其中acc過表達(dá)的酵母細(xì)胞溶解產(chǎn)物，但是與對照相比沒有檢測到吸光度變化。放射性acc酶測定是非常敏感的，甚至可以檢測pmol/mg/min活性。首先用市售的純化的人acc酶測試該方法。當(dāng)檢測到明顯的活性時，進(jìn)一步優(yōu)化該方法以檢測酵母細(xì)胞溶解產(chǎn)物的acc活性。測定表13中列出的菌株的acc1活性，并基于該結(jié)果制作描述acc1的相對排名的列表(圖2)。對最有前途的候選者accyl進(jìn)行更多的研究，釀酒酵母s-128野生型菌株的結(jié)果和其中acc1已被過表達(dá)的相同菌株的結(jié)果顯示在表14中。表14acc1基因的過表達(dá)導(dǎo)致acc1酶活性是野生型釀酒酵母vsk-128菌株的內(nèi)源acc1活性的2.9倍。aadh體外酶活性測定最初選擇了5種aadh基因(即adhepm、adheec、eutec、eutdz和lin1129li)，并轉(zhuǎn)化到cen.pk實(shí)驗(yàn)室菌株中。這些aadh從多拷貝質(zhì)粒表達(dá)，其中它們在tef1啟動子的控制下。所有五個aadh基因顯示aadh活性，但是與adhe型aadh(即adhepm和adheec)相比，三種eute型aadh基因(即eutec、eutdz和lin1129li)在酵母中給出高得多的aadh活性。從基因組挖掘分析中選擇20個其他新aadh基因以評估酵母中的表達(dá)和酶活性。測試這二十個新的aadh基因的體外酶活性，其中四個(即aadhmm、aadhab、aadhbw和aadhvs)顯示具有aadh活性。aadh體內(nèi)生長測定從工業(yè)釀酒酵母vsk-128菌株中刪除所有acs2基因拷貝以產(chǎn)生具有缺陷型pdh旁路的菌株，該菌株在基于葡萄糖的培養(yǎng)基上不能生長。然后將攜帶25種不同aadh變體的表達(dá)載體轉(zhuǎn)化到該acs2缺失菌株中，以評價它們恢復(fù)該菌株在葡萄糖上的生長的能力?；诰暝谄咸烟腔傊桨迳仙L的能力，12種不同的aadh變體(即euteec、eutedz、lin1129li、aadhmm、aadht1、aadhab、aadhta、aadhbs、aadhbw、aadhvs、aadhhs和adheec)能夠恢復(fù)acs2-缺失菌株的生長。體外aadh酶活性測定和體內(nèi)生長恢復(fù)分析之間存在良好的相關(guān)性。然后在液體搖瓶培養(yǎng)中評估被發(fā)現(xiàn)能夠恢復(fù)acs2缺失菌株生長的前九個aadh變體菌株的生長速率，并與含有完整pdh旁路的野生型菌株進(jìn)行比較。當(dāng)在葡萄糖上生長時，所有9個這些aadh變體都能夠維持>50％的釀酒酵母vsk-128菌株有氧生長速率，并且這些aadh變體中的6個能夠維持≥80％的釀酒酵母vsk-128菌株有氧生長速率。酵母中的mcr酶測定將八個全長綠屈撓菌屬mcr同源物截短成它們的兩個功能結(jié)構(gòu)域，并且測定mcr特異性結(jié)構(gòu)域以及六個古細(xì)菌mcr在酵母中的酶活性。這些mcr從多拷貝質(zhì)粒表達(dá)，其中它們在tef1啟動子的控制下并轉(zhuǎn)化到cen.pk實(shí)驗(yàn)室菌株中。當(dāng)利用nadph作為輔因子時，mcrca(橙色綠屈撓菌)及其同源物mcrrc(光合玫瑰彎菌)是僅有的兩種給出比野生型菌株更高的mcr酶活性的綠屈撓菌屬mcr同源物。當(dāng)利用nadh作為輔因子時，沒有觀察到這八種綠屈撓菌屬mcr同源物的酶活性。當(dāng)利用nadph作為輔因子時，三個古細(xì)菌mcr[sulfolobalesarchaeon、嗜酸熱硫化葉菌(x2)]給出了比野生型菌株高的mcr酶活性，使用nadh作為輔因子時，所有六個古細(xì)菌mcr都給出比野生型菌株高的mcr酶活性。古細(xì)菌mcr的異源表達(dá)和表征為四種不同的古細(xì)菌mcr(勤奮金屬球菌(mcrms)、頭寇岱硫化葉菌(mcrst)，candidatuscaldiarchaeum(mcrcc)和古菌硫化葉菌(mcrsa1))制備基于pbatt7啟動子的表達(dá)構(gòu)建體。這些構(gòu)建體不含有純化標(biāo)簽，是大腸桿菌密碼子優(yōu)化的且在本報告前面所述的條件下表達(dá)(表15)。表15用于在大腸桿菌中表達(dá)的古細(xì)菌mcr構(gòu)建體的概述(ca-橙色綠屈撓菌，ms-勤奮金屬球菌，st-頭寇岱硫化葉菌，cc-candidatuscaldiarchaeum，sa1-古菌硫化葉菌，sa2-嗜酸熱硫化葉菌)使用以下測定條件分析它們的活性：0.4mmnad(p)h，0.15mm丙二酰輔酶a，tris-hclph7，2mmmgcl2。在室溫下以微量滴定板(mtp)形式對20倍稀釋的溶解產(chǎn)物進(jìn)行測定。隨后及時追蹤a365處的nadph或nadh氧化。在測試的四個古細(xì)菌基因中，mcrsa1和mcrst顯示最高的mcr活性。然而，mcr活性小于測量的標(biāo)記的mcrca的活性。對于candidatuscaldiarchaeummcr(mcrcc)，沒有測量到對nadph或nadh的活性(在大腸桿菌細(xì)胞溶解產(chǎn)物中)。當(dāng)使用sds-page凝膠分析這些構(gòu)建體時，mcrsa1在大腸桿菌溶解產(chǎn)物中顯示最高的表達(dá)水平，而mcrcc不能以可溶形式表達(dá)；參見凝膠(sa1>ca>st>ms>cc)(圖3)。構(gòu)建體mcrsa1和mcrst似乎具有雙重輔因子偏好，并且與測量到的對nadph的活性相比，顯示出約40-50％的相對nadh活性。就比活性而言，mcrst可以是所測試的四種古細(xì)菌mcr中活性最高的酶，因?yàn)樗谙鄬Φ偷谋磉_(dá)水平下顯示相對高的活性水平。實(shí)施例4通過培養(yǎng)重組酵母產(chǎn)生3-hp釀酒酵母的搖瓶培養(yǎng)從在選擇性瓊脂基平板上新生長的菌株中取出一小圈細(xì)胞，用于接種在250ml燒瓶中的20ml選擇性sc基培養(yǎng)基(20g/l葡萄糖)，并生長2天(30℃，250rpm)，直到所有葡萄糖和乙醇已被消耗。測定最終細(xì)胞密度，將培養(yǎng)物在4000rpm離心5分鐘。然后通過hplc或gc/ms分析上清液以確定培養(yǎng)物上清液中3-hp和其它主要代謝物的積累。然而，還根據(jù)具體菌株和目的來測試其他培養(yǎng)條件(例如葡萄糖的起始量、通氣量、培養(yǎng)基的類型和向培養(yǎng)基中加入另外的物質(zhì)等)。實(shí)施例5釀酒酵母的生物反應(yīng)器培養(yǎng)在含有250-500ml培養(yǎng)基的multifors生物反應(yīng)器(最大工作體積500ml，24葉片rusthon渦輪葉輪，inforsht，瑞士)中進(jìn)行培養(yǎng)。最初將培養(yǎng)物維持在30℃，300或900-950rpm，1.2、2.4或3.6體積氣體(體積培養(yǎng))-1min-1(vvm)。通過加入無菌2mnaoh或2mh3po4使培養(yǎng)物ph在ph5.5±0.2處恒定。加入clerolfba3107消泡劑(cognisfrance，ponthierryparis；0.03％v/v)以控制泡沫產(chǎn)生。在用含3％co2的ar；含5％co2、0.99％ar、15％o2的n2；含20％o2加20％ar的n2；含0.04％乙醇的n2校準(zhǔn)的primaproprocess質(zhì)譜儀(thermoscientific，英國)中連續(xù)分析氣體濃度(co2、o2、n2和ar)。將菌株在搖動的燒瓶中在scd基選擇性培養(yǎng)基中預(yù)生長過夜，并用于接種生物反應(yīng)器。使分批培養(yǎng)階段(20g/l初始葡萄糖)持續(xù)14至20小時，并且僅在葡萄糖已被消耗后開始葡萄糖進(jìn)料，但葡萄糖進(jìn)料在乙醇已被消耗之前或之后開始(取決于培養(yǎng)目的)。葡萄糖進(jìn)料速率保持在0.38-0.65gl-1h-1(取決于培養(yǎng)目的)。然后通過hplc分析上清液樣品，以確定培養(yǎng)物中3-hp和其他主要代謝物的積累。實(shí)施例6通過hplc對細(xì)胞培養(yǎng)物上清液進(jìn)行3-hp分析用watersalliancee2695hplc系統(tǒng)(waters，美國米爾福德)分析培養(yǎng)物上清液樣品，其中注射體積為10μl。使用與fastacidanalysiscolumn(100mm×7.8mm)(bio-rad，美國)連接的aminexhpx-87horganicacidcolumn(300mm×7.8mm)(bio-rad，美國)作為hplc的固定相。將柱保持在+55℃，并使用5.0mmh2so4(merckkgaa，德國)作為洗脫液，流速為0.3mlmin-1或0.5mlmin-1。使用waters2489雙波長uv(210nm)檢測器(waters，美國米爾福德)和waters2414差示折射計(waters，美國米爾福德)檢測3-羥基丙酸、葡萄糖、乙酸、琥珀酸、丙酮酸、甘油和乙醇。實(shí)施例7通過gc/ms對細(xì)胞培養(yǎng)物上清液進(jìn)行3-hp分析按以下方式制備測試樣品并制作標(biāo)準(zhǔn)曲線：用50μlhcl(6n)酸化上清液(0.5ml)，并摻加3-hpa(tci)標(biāo)準(zhǔn)品(在乙酸乙酯中)。加入5μl的乳酸內(nèi)標(biāo)溶液(sigmaaldrich(isotec)l-乳酸鈉-3,3,3-d398原子％；5.5g/l)和約0.2g的nacl。由于標(biāo)記的3-hp不可商購獲得，因此選擇該乳酸穩(wěn)定同位素產(chǎn)品作為內(nèi)標(biāo)，因?yàn)樵摦a(chǎn)品是可獲得的、在樣品基質(zhì)中不存在、與3-hp在結(jié)構(gòu)/化學(xué)方面最相似的化合物。將混合物在渦旋混合器中振蕩約3分鐘。然后采用0.5ml乙酸乙酯通過在渦旋混合器中混合約3分鐘萃取樣品兩次。通過在10000rpm離心5分鐘分離各層。將上層收集到gc小瓶中并蒸發(fā)。通過在60℃下孵育1小時，用含有1％tmcs的mstfa(50μl)將干燥的殘余物衍生化。以與樣品相同的方式提取校準(zhǔn)曲線的標(biāo)準(zhǔn)品以使誤差最小化。在與agilent5973質(zhì)量選擇性檢測器(msd)結(jié)合的agilent6890氣相色譜儀(gc)上運(yùn)行樣品。注射器(注射體積1μl，分流比20:1)和msd界面溫度為280℃，并且爐溫程序?yàn)橐?0℃/min的速率從50℃升至280℃。在agilenthp-5ms毛細(xì)管柱(30m，id200μm，膜厚度0.25μm；agilent19091s-433)上進(jìn)行分析?；衔锏蔫b定基于對nist文庫的光譜搜索。通過監(jiān)測m/z147和m/z219檢測3-hp，通過監(jiān)測m/z177檢測3-hp二聚體。通過利用3-hp響應(yīng)(responses)構(gòu)建5點(diǎn)校準(zhǔn)曲線(c＝1-400mg/l)，并使用內(nèi)標(biāo)歸一化。證明在該濃度范圍內(nèi)的定量是線性的?？瞻讟悠放c樣品一起分析。實(shí)施例8釀酒酵母質(zhì)粒表達(dá)菌株的評價3-hp質(zhì)粒表達(dá)菌株同時從兩種不同的表達(dá)質(zhì)粒(即psk-084和/或psk-085)表達(dá)一種、兩種或三種3-hp途徑酶(即aadh、acc1、mcr和hpdh)(圖4)。這些菌株用于評價不同的3-hp途徑酶(和這些酶的組合)對vsk-128耐酸釀酒酵母菌株中3-hp的產(chǎn)量的影響。將菌株在250ml燒瓶中的20ml選擇性sc基培養(yǎng)基(20g/l葡萄糖)中培養(yǎng)，并生長2天(30℃，250rpm)直到所有葡萄糖和乙醇被消耗掉。實(shí)施例9體內(nèi)途徑酶活性分析的總結(jié)對表16中的25個aadh、8個acc1、10個雙功能hpdh-mcr、6個古細(xì)菌mcr和28個hpdh在各種釀酒酵母菌株中產(chǎn)生3-hp的能力進(jìn)行分析，如果需要，所述釀酒酵母菌株也表達(dá)額外的3-hp途徑酶。顯示許多新的3-hp途徑酶(從基因組挖掘分析獲得)在酵母中是有活性的，并且與先前公開的3-hp途徑酶相比，它們中的許多具有更優(yōu)異的性質(zhì)(即更高的活性，更好的輔因子偏好)。表16實(shí)施例10用于3-hp生產(chǎn)的釀酒酵母搖瓶培養(yǎng)試驗(yàn)培養(yǎng)條件在一對早期測試菌株上評價各種不同的培養(yǎng)條件，以觀察各種培養(yǎng)條件如何影響菌株產(chǎn)生3-hp的能力。釀酒酵母vsk-128(δura3，δhis3)菌株表達(dá)兩種質(zhì)粒，其中一種質(zhì)粒含有hibadhpa或hpdhec基因，第二種質(zhì)粒含有mcrsa2基因。兩種菌株在培養(yǎng)期間表現(xiàn)相似，并且具有非常相似的代謝物譜圖。測試六種不同的搖瓶培養(yǎng)條件：1.有氧，分批法，高初始葡萄糖(120g/l)。在第3天再加入100g/l葡萄糖。2.無氧，分批法，高初始葡萄糖(100g/l)。將燒瓶密封并且搖動更慢，速度為100rpm。3.有氧，分批法，重復(fù)摻加葡萄糖(glucosespiking)。初始葡萄糖為20g/l，然后每隔一天加入40g/l。4.有氧，模擬分批補(bǔ)料法，許多初始葡萄糖片。最初加入5片，在第3天再加入5片。每天加入不同量的酶a溶液(50-150μl/天)。5.有氧，模擬分批補(bǔ)料法，重復(fù)摻加葡萄糖片。最初加入1片，在第1天和第2天加入2片，在第3天和第4天加入3片。每天加入不同量的酶a溶液(50-150μl/天)。6.好氧，分批法，重復(fù)摻加半乳糖。初始半乳糖為20g/l，然后每隔一天加入40g/l。對于模擬補(bǔ)料分批培養(yǎng)，每個片劑被認(rèn)為釋放0.5g葡萄糖(對于25-ml培養(yǎng)體積而言，其相當(dāng)于20g/l葡萄糖)。片劑基本上由葡萄糖(即淀粉)組成，酶a溶液(即淀粉酶)允許在搖瓶培養(yǎng)期間將葡萄糖受控地緩慢釋放到培養(yǎng)基中。假設(shè)葡萄糖受限的補(bǔ)料分批條件可以促進(jìn)生長和隨后的3-hp生產(chǎn)的效率(flux)。細(xì)胞生長所有正常的基于葡萄糖的培養(yǎng)均類似地生長，并且半乳糖補(bǔ)料培養(yǎng)生長更慢。另一方面，與其他培養(yǎng)條件相比，分批補(bǔ)料條件促進(jìn)更多的細(xì)胞生長。特別地，分批補(bǔ)料(片劑摻加)條件在培養(yǎng)早期的確促進(jìn)了大量生長。3-hp生產(chǎn)對于這些培養(yǎng)，大多數(shù)生長通常在第2天結(jié)束時發(fā)生，并且絕大多數(shù)3-hp也在第2天生成，因此表明生長和3-hp產(chǎn)生彼此相關(guān)。分批補(bǔ)料(片劑摻加)培養(yǎng)條件產(chǎn)生最多的3-hp(～0.85g/l)，半乳糖進(jìn)料培養(yǎng)產(chǎn)生最少量的3-hp(～0.12g/l)，其他培養(yǎng)條件全部產(chǎn)生約0.3-0.4g/l的3-hp。這些結(jié)果表明培養(yǎng)條件可對3-hp產(chǎn)生水平有大的影響(圖5)。葡萄糖消耗在所有培養(yǎng)試驗(yàn)中葡萄糖被快速消耗直到第2-3天，然后葡萄糖消耗速率隨著生長和3-hp產(chǎn)生顯著降低(在第4-5天期間)。對于補(bǔ)料分批培養(yǎng)，似乎葡萄糖的消耗與其從片劑釋放一樣快，表明這些培養(yǎng)在葡萄糖受限的條件下進(jìn)行。甘油、乙酸和乙醇積累對于葡萄糖進(jìn)料培養(yǎng)，甘油積累是相當(dāng)高的，對于半乳糖進(jìn)料培養(yǎng)而言甘油積累則低得多。乙酸積累在不同的培養(yǎng)條件中相當(dāng)相似，但是補(bǔ)料分批(許多片劑)條件產(chǎn)生更多的乙酸。對于這些搖瓶培養(yǎng)中的大部分，特別是對于葡萄糖分批培養(yǎng)和葡萄糖摻加培養(yǎng)，存在大量的乙醇累積。補(bǔ)料分批培養(yǎng)旨在代表葡萄糖受限條件以減少過量溢流代謝生成乙醇，并且這種方法似乎已經(jīng)取得成功，因?yàn)樵谘a(bǔ)料分批培養(yǎng)中乙醇積累顯著較低。培養(yǎng)更成熟的3-hp產(chǎn)生菌株按照最有前途的補(bǔ)料分批(片劑摻加)培養(yǎng)條件培養(yǎng)其他3-hp生產(chǎn)菌株[(euteec、hpdh-mcrca、acc1sc_s1157a)和(hibadhpa、mcrsa2)]以檢查其性能。同樣，大部分生長發(fā)生在第3天，大部分3-hp產(chǎn)生發(fā)生在第3天。在該培養(yǎng)條件下，該菌株的3-hp產(chǎn)量超過1.2g/l(圖6)。盡管詳細(xì)描述了本發(fā)明的具體實(shí)施方案，但是對于本領(lǐng)域技術(shù)人員來說顯而易見的是，具體描述僅僅是令人滿意的示例性實(shí)施方案，并且不應(yīng)被解釋為限制本發(fā)明的范圍。因此，本發(fā)明的實(shí)質(zhì)范圍由所附權(quán)利要求及其等同物限定。工業(yè)適用性使用本發(fā)明的重組酵母和制備3-hp的方法使得能夠以高濃度和高收率在低ph下從有用的糖(如葡萄糖)生產(chǎn)3-hp，從而極大地有助于從生物質(zhì)經(jīng)濟(jì)地生產(chǎn)3-hp及其應(yīng)用產(chǎn)品。序列表<110>sk新技術(shù)株式會社<120>產(chǎn)生3-羥基丙酸的重組酵母和使用其產(chǎn)生3-羥基丙酸的方法<130>pf-b1841<140>pct/kr2015/009061<141>2015-08-28<150>10-2014-0114505<151>2014-08-29<160>144<170>patentinversion3.2<210>1<211>2248<212>prt<213>kazachstaniaexigua<400>1metserglugluasnleuphegluaspserseralalysargglntyr151015gluilethrasptyrserlyslyshislysalaleualaprohisphe202530valglyleuasnthrleuasplysvalglugluserproleulysasp354045phevallysserhisglyglyhisthrvalileserlysileleuile505560alaasnasnglyilealaalavallysgluileargservalarglys65707580trpsertyrgluthrpheglyaspgluargthrvalglnphevalala859095metalathrprogluaspleuglnalaasnserglutyrileargmet100105110alaaspglntyrvalgluvalproglyglythrasnasnasnasntyr115120125alaasnvalaspleuilevalaspvalalagluargthraspvalasp130135140alavaltrpalaglytrpglyhisalasergluasnprohisleupro145150155160glulysleualaalaserlysarglysileilepheileglypropro165170175glyseralametargserleuglyasplysileserserthrileval180185190alaglnhisalalysvalprocysileprotrpserglythrglyile195200205asplysvalhisvalaspgluthrserglyleuvalservalaspasp210215220aspvaltyrglnglnglycyscysglnserprogluaspglyleuala225230235240lysalalysglnileglypheprovalmetilelysalaserglugly245250255glyglyglylysglyileargglnvalgluargglugluasppheile260265270alaleutyrhisglnalaalaasngluileproglyserproilephe275280285ilemetlysleualaglylysalaarghisleugluvalglnleuleu290295300alaaspglntyrglythrasnileserleupheglyargaspcysser305310315320valglnargarghisglnlysileilegluglualaprovalthrile325330335alaasnasngluthrpheasplysmetalaasnalaalavalargleu340345350glylysleuvalglytyrvalseralaglythrvalglutyrleutyr355360365serasngluglulyslystyrtyrpheleugluleuasnproargleu370375380glnvalgluhisprothrthrglumetvalserglyvalasnleupro385390395400seralaglnleuglnilealametglyilepromethisargileser405410415aspileargvalleutyrglymetasnprohisseralathrgluile420425430aspphegluphelysthrgluglualalyslysthrglnarglyspro435440445valprolysglyhiscysthralacysargilethrsergluasppro450455460asngluglyphelysproserglyglyserleuhisgluleuasnphe465470475480argserserserasnvaltrpglytyrpheservalglyasnasngly485490495glyilehisserpheseraspserglnpheglyhisilephealaphe500505510glygluasnargglnmetserarglyshismetvalvalalaleulys515520525gluleuserileargglyasppheargthrthrvalglutyrleuile530535540lysleuleugluthrgluasppheglugluasnthrilethrthrgly545550555560trpleuaspaspleuileserhislysmetthralagl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