CN114615881B - 产生甘油三酯的非人类生物体 - Google Patents
产生甘油三酯的非人类生物体 Download PDFInfo
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Abstract
本文描述了用于产生甘油三酯的非人类生物体，其中所述非人类生物体经遗传修饰表达对C16：0‑辅酶A具有特异性的溶血磷脂酸酰基转移酶(C16：0LPAT)，其中，一旦表达，C16：0LPAT位于内质网中。还描述了用于产生甘油三酯的非人类生物体，其中(a)对C16：0‑辅酶A(C16：0LPAT)具有特异性的溶血磷脂酸酰基转移酶位于内质网中；(b)抑制或阻止天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。进一步描述了在其中表达叶绿体溶血磷脂酸酰基转移酶(LPAT)的非人类生物体，其中所述叶绿体LPAT缺乏功能性叶绿体靶向信号。本发明还涉及从生物体中提取甘油三酯的方法及其在婴儿配方奶粉中的用途。
Description
技术领域
本发明涉及用于产生甘油三酯的非人类生物体，特别是植物、真菌和酵母。
背景技术
婴儿配方奶粉(infant formula)是一种替代人乳的人造食品。人乳中大约一半的卡路里由脂肪(甘油三酯；TAG)提供，而在婴儿配方奶粉中，这种脂肪主要来自植物。虽然混合植物脂肪可以复制主要包含棕榈酸酯(C16：0)和油酸酯(C18：1)的人乳脂(HMF)的脂肪酰基组成，但其酯化骨架在甘油的酰基布置(即立体异构结构)与人乳完全不同。在植物脂肪中，饱和长链脂肪酰基(例如C16：0)占据外侧立体专一编号(sn)位置(sn-1/3)，几乎被排除在中间(sn-2或β)位置之外。而在HMF中，70％以上的C16：0位于sn-2位，不饱和脂肪酰基(主要是C18：1)占据外部sn-1/3位。
多项针对早产儿和足月儿的临床试验表明，HMF这种不寻常的立体异构结构对新生儿肠道中的营养吸收非常重要。提出了如下机制。在肠道消化阶段，脂肪酶攻击摄入的sn-1/3位的脂肪，产生易于吸收的2-单酰基甘油。当不饱和脂肪酸从sn-1/3位释放时，它们也很容易被吸收，但长链饱和脂肪酸(例如C16：0)的释放存在问题。它们的熔点高于体温，在肠道pH值下，它们很容易与钙和镁等矿物质形成水合脂肪酸皂类。因此，C16：0分布在植物脂肪sn-1/3位意味着它们比HMF更难吸收。有证据表明，C16：0皂类的形成还会降低钙的吸收，从而损害早期骨骼发育，并且这些皂类在肠道中的积聚还会干扰输送，导致婴儿不适。
为了模仿HMF的立体异构结构，几家公司开发了HMF替代品(HMFS)。HMFS是利用三棕榈酸甘油酯、不饱和游离脂肪酸(主要是C18：1)和固定化重组sn-1/3-区域选择性脂肪酶，通过酶催化酸解(或醇解和酯化)制备的。HMFS的价格远高于传统植物脂肪混合物的价格，主要是酶催化增加了成本，包括产生了有机溶剂废物。还可以提供不同牌号的HMFS，提供在sn-2位具有～40％至～70％的C16：0的全脂阶段。真正的HMF模拟物(>70％的C16：0在sn-2位)生产成本最高，因为它们需要采用两步催化工艺和通过特殊分馏程序和化学随机化从棕榈油得到的三棕榈酸甘油酯纯原料。价格和质量之间的平衡可能是限制HMFS使用的一个因素，尽管越来越多的临床证据表明这种成分是有益的，但目前仅在大约10％的婴儿配方奶粉中使用，特别是在为易于消化而配制和销售的高端产品中使用。即使在这些产品中，C16：0在sn-2位的富集与HMF相比仍然存在很大差距。
因此，本发明的目的是寻求缓解上述问题。
发明内容
根据本发明的一个方面，提供用于产生甘油三酯的非人类生物体，其中所述非人类生物体经遗传修饰表达对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶(C16：0LPAT)，其中，一旦表达，C16：0LPAT定位于内质网。
优选地，天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性在生物体中受到抑制或阻止。
根据本发明的另一方面，提供用于产生甘油三酯的非人类生物体，其中：-
(a)对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶(C16：0LPAT)位于内质网中；和
(b)天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性受到抑制或阻止。
优选地，二酰基甘油转化为磷脂酰胆碱并且从磷脂酰胆碱转化的活性受到抑制或阻止。
优选地，一种或多种天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性受到抑制或阻止。
根据本发明的另一方面，提供用于产生甘油三酯的非人类生物体，其中：-
(a)对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶(C16：0LPAT)位于内质网中；
(b)天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性受到抑制或阻止；和
(c)二酰基甘油转化为磷脂酰胆碱并且从磷脂酰胆碱转化的活性受到抑制或阻止。
值得注意的是，正如本文进一步详细讨论的那样，通过提供位于内质网中对C16：0-辅酶A(CoA)具有特异性的溶血磷脂酸酰基转移酶，其在胞质甘油脂生物合成途径中发挥作用，在TAG生物合成过程中将C16：0酯化到中间位置。这种情况与天然内质网溶血磷脂酸酰基转移酶的抑制和任选二酰基甘油与磷脂酰胆碱的相互转化的抑制相结合，意味着与外部位置相比，更多的C16：0被酯化到甘油骨架的中间位置。
优选地，生物体包含内质网。
优选地，生物体是植物、真菌、酵母或藻类。
优选地，生物体不是哺乳动物。
优选地，生物体是植物。优选地，植物是油料植物。
优选地，生物体是属于十字花目、菊目、豆目、金虎尾目、锦葵目、蔷薇目、唇形目、茄目、槟榔目或禾本目的植物。最优选地，植物属于十字花目或菊目，更优选十字花目。
优选地，生物体是属于十字花科、菊科、豆科、亚麻科、锦葵科、大麻科、胡麻科、木犀科、茄科、槟榔科或禾本科的植物。最优选地，植物属于十字花科或菊科，更优选十字花科。
优选地，生物体是属于拟南芥属、亚麻荠属、芸苔属、菥蓂属、独行菜属、向日葵属、大豆属、花生属、亚麻属、烟草属、棉属、大麻属、胡麻属、木犀榄属、油棕属、玉蜀黍属、燕麦属或稻属的植物。最优选地，植物属于拟南芥属、亚麻荠属、芸苔属或向日葵属。
优选地，植物选自拟南芥、亚麻荠、欧洲油菜、埃塞俄比亚芥、甘蓝、芜菁、芥菜、菥蓂、家独行菜(Lepidium sativum)、向日葵、大豆、花生、亚麻、陆地棉、大麻、芝麻、油橄榄、本氏烟(Nicotiana benthamiana)、油棕、美洲油棕、玉蜀黍、燕麦或稻。
优选地，所述植物选自拟南芥(Arabidopsis thaliana)、向日葵(Helianthusannuus)、大豆(Glycine max)、亚麻荠(Camelina sativa)和欧洲油菜(Brassica napus)。
优选地，所述植物选自拟南芥、亚麻荠和欧洲油菜。
优选地，所述植物是拟南芥。
优选地，所述生物体是可以从例如植物的种子、果实或叶子中提取甘油三酯的植物。
优选地，油料植物是可以从种子或果实中提取甘油三酯的植物。
优选地，生物体是酵母。
优选地，生物体是发酵酵母和/或呼吸酵母。
优选地，生物体是产油酵母。
优选地，生物体是能够在细胞生物质中积累至少约20％甘油三酯的酵母。
优选地，生物体是属于酵母目、酵母菌目、银耳目、黑粉菌目、锁掷酵母目、毛霉菌目、被孢霉目或散囊菌目的真菌。最优选生物体来自酵母目。
优选地，生物体是属于双足囊菌科(Dipodascaceae)、酵母科、复膜孢酵母科、银耳科、黑粉菌科、锁掷酵母科(Sporidiobolaceae)、丝孢酵母科(Trichosporonaceae)、法夫酵母科(Phaffomycetaceae)、毛红曲霉科(Trichomonascaceae)、毛霉菌科、被孢霉科、小克银汉霉科或发菌科的真菌。最优选生物体来自双足囊菌科。
优选地，生物体是属于酵母属、耶氏酵母属、隐球菌属、念珠菌属、红冬孢酵母属、红酵母属、油脂酵母属、毛孢子菌属、威克汉姆酵母属、毕赤酵母属、拟内孢霉属、接合囊酵母属、毛霉属、被孢霉属、小克银汉霉属或曲霉属的真菌。最优选生物体来自耶氏酵母属。
优选地，生物体是专性呼吸酵母和克拉布特里阴性(Crabtree-negative)酵母。
优选地，生物体是解脂耶氏酵母(Yarrowia lipolytica)。
优选地，生物体是解脂耶氏酵母，C16：0LPAT是CreLPAT。
优选地，生物体是解脂耶氏酵母的gsy1Δ-CreLPAT菌株。
优选地，生物体在包含碳源的培养基中培养，其中碳源包含一种或多种糖。
优选地，一种或多种糖包括一种或多种可发酵糖。
优选地，一种或多种糖选自木糖、乳糖、纤维素、葡萄糖、果糖、蔗糖或水解的木质纤维素材料中的一种或多种。
优选地，生物体在包含碳源的培养基中培养，其中碳源包含一种或多种脂肪酸和/或脂肪酸酯。
优选地，碳源包含C16：0。
优选地，碳源包含脂肪酸和/或脂肪酸酯的混合物，其中至少约30％w/w的脂肪酸和/或脂肪酸酯包含C16：0和/或至少约30％w/w的脂肪酸和/或脂肪酸酯包含C18：1。
优选地，生物体在包含碳源的培养基中培养，其中碳源包含一种或多种植物油。
优选地，生物体在包含碳源的培养基中培养，其中碳源包含(i)一种或多种糖和(ii)一种或多种脂肪酸和/或脂肪酸酯的混合物。
优选地，生物体在包含碳源的培养基中培养，其中碳源包含(i)一种或多种糖和(ii)一种或多种油，优选植物油的混合物。
优选地，碳源包含甘油。
优选地，生物体在包含碳源的培养基中培养，其中碳源选自甘油、葡萄糖和/或棕榈油。
优选地，碳源包含棕榈油。
优选地，碳源包含棕榈油与葡萄糖和/或甘油的组合。
优选地，碳源的浓度是至少约5gL-1，优选至少约10gL-1，优选至少约15gL-1，优选至少约20gL-1。
优选地，碳源的浓度是约5gL-1至约35gL-1，优选约10gL-1至约30gL-1，优选约15gL-1至约25gL-1，优选浓度是约20gL-1。
优选地，培养基包含浓度至少约2.5gL-1的第一碳源和浓度至少约2.5gL-1的第二碳源。
优选地，培养基包含浓度至少约5gL-1，优选至少约10gL-1的第一碳源，和浓度至少约5gL-1，优选至少约10gL-1的第二碳源。
优选地，培养基包含浓度约2.5gL-1至约20gL-1的第一碳源，和浓度约2.5gL-1至约20gL-1的第二碳源。
优选地，培养基包含浓度约5gL-1至约15gL-1，优选约10gL-1的第一碳源，和浓度约5gL-1至约15gL-1，优选约10gL-1的第二碳源。
优选地，第一碳源包含一种或多种糖。
优选地，一种或多种糖包括一种或多种可发酵糖。
优选地，一种或多种糖选自木糖、乳糖、纤维素、葡萄糖、果糖、蔗糖或水解的木质纤维素材料中的一种或多种。
优选地，第一碳源包含甘油和/或一种或多种糖。
优选地，第二碳源包含一种或多种脂肪酸和/或脂肪酸酯。
优选地，第二碳源包含C16：0。
优选地，第二碳源包含脂肪酸和/或脂肪酸酯的混合物，其中至少约30％w/w的脂肪酸和/或脂肪酸酯包含C16：0和/或至少约30％w/w的脂肪酸和/或脂肪酸酯包含C18：1。
优选地，第二碳源包含一种或多种植物油。
优选地，第二碳源包含棕榈油。
优选地，培养基是限氮培养基。
优选地，培养基包含碳和氮，其中碳与氮之比大于约40。
优选地，生物体是酵母，C16：0LPAT是AGPAT1。
优选地，生物体是酿酒酵母，C16：0LPAT是AGPAT1。
根据本发明的另一个方面，提供非人类生物体，其中所述生物体是经遗传修饰表达AGPAT1的酿酒酵母。
优选地，生物体是无绿藻(Prototheca algae)。
优选地，C16：0LPAT选自：-
(i)缺乏功能性叶绿体靶向序列的叶绿体LPAT，优选地，缺乏功能性叶绿体靶向序列的植物叶绿体LPAT；
(ii)AGPAT1(人类1-酰基甘油-3-磷酸-O-酰基转移酶亚型1)；
(iii)CreLPAT；
(iv)微拟球藻属(Nannochloropsis sp.)LPAT2；
(v)微拟球藻属LPAT3；
(vi)微拟球藻属LPAT4；和
(vii)集胞藻属(Synechocystis sp.)LPAT。
优选地，生物体经处理或遗传修饰破坏叶绿体靶向序列。
优选地，叶绿体LPAT经处理或遗传修饰破坏叶绿体靶向序列。
优选地，叶绿体LPAT是缺少至少一部分、优选全部叶绿体靶向序列的修饰叶绿体LPAT，优选是缺少至少一部分、优选全部叶绿体靶向序列的修饰植物叶绿体LPAT。
优选地，叶绿体靶向序列是N-末端叶绿体靶向序列。
优选地，生物体经修饰或处理(例如通过遗传修饰，优选通过叶绿体靶向序列的遗传修饰)破坏叶绿体LPAT叶绿体靶向序列。
优选地，所述遗传修饰包括添加或删除核酸和/或氨基酸残基。
优选地，生物体经遗传修饰表达位于内质网中对C16：0-CoA(C16：0LPAT)具有特异性的溶血磷脂酸酰基转移酶。
优选地，生物体是植物，C16：0LPAT在种子特异性启动子，优选油质蛋白、油菜籽蛋白或大豆球蛋白种子特异性启动子，最优选种子特异性大豆球蛋白-1启动子(ProGLY)的控制下表达。
优选地，生物体经处理或遗传修饰以抑制或阻止天然ER LPAT的活性。
优选地，生物体经处理或遗传修饰以抑制或阻止一种或多种天然ER LPAT的活性。
优选地，ER LPAT对C16：0-CoA不具有特异性。
优选地，使用人工微RNA抑制或阻止天然ER LPAT的活性。
优选地，使用RNAi和/或基因组编辑和/或突变育种来抑制或阻止天然ER LPAT的活性。
优选地，人工微RNA由种子特异性启动子(优选油质蛋白、油菜籽蛋白或大豆球蛋白种子特异性启动子)控制。
优选地，生物体在ER LPAT翻译起始位点的非编码区5'中包含破坏性插入。优选地，破坏性插入位于ER LPAT翻译起始位点的约139bp 5'。优选地，破坏性插入是ER LPAT翻译起始位点的139bp 5'。优选地，破坏性插入是T-DNA插入。
或者，破坏性插入位于ER LPAT翻译起始位点的约302bp 5'，例如302bp 5'。
优选地，生物体中二酰基甘油向磷脂酰胆碱的转化受到抑制或阻止。
优选地，通过抑制或阻止胆碱磷酸转移酶(CPT1)和/或乙醇胺磷酸转移酶(EPT1)的活性来抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化。
优选地，生物体经处理或遗传修饰破坏CPT1和/或EPT1。
优选地，抑制或阻止生物体中磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性。
优选地，抑制或阻止生物体中一种或多种磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性。
优选地，通过抑制或阻止生物体中磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性，例如通过抑制或阻止生物体中一种或多种磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性来降低二酰基甘油和磷脂酰胆碱的相互转化。
优选地，生物体经处理或遗传修饰增加总C16：0的含量。
优选地，生物体是经处理或遗传修饰增加优选其种子、果实或叶子中，最优选种子或果实中总C16：0含量的植物。
优选地，生物体表达FATB硫酯酶，优选拟南芥FATB硫酯酶。
优选地，生物体经遗传修饰表达FATB硫酯酶。
优选地，生物体经处理或遗传修饰以破坏质体β-酮脂酰-ACP合酶II。
优选地，生物体经处理或遗传修饰以破坏KASII基因FAB1(At1g74960)。
优选地，生物体经处理或遗传修饰以破坏脂肪酸延长酶基因(FAE1)。
优选地，生物体经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)。
优选地，生物体表达FATB硫酯酶，优选拟南芥FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)。
优选地，生物体经处理或遗传修饰表达FATB硫酯酶基因，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)。
优选地，生物体，优选植物，经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)。
优选地，(i)生物体，优选植物，经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ERLPAT)的活性。
优选地，(i)生物体，优选植物，经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ERLPAT)的活性，及(iv)C16：0LPAT是ΔCTS-LPAT1或AGPAT1。
优选地，(i)生物体，优选植物，表达FATB硫酯酶，优选拟南芥FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
优选地，(i)生物体，优选植物，表达FATB硫酯酶，优选拟南芥FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性，及(iv)C16：0LPAT是ΔCTS-LPAT1或AGPAT1。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达C16：0LPAT，并且其中(i)生物体经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达C16：0LPAT，并且其中(i)生物体表达FATB硫酯酶，优选拟南芥FATB硫酯酶，经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达ΔCTS-LPAT1，并且其中(i)生物体经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达ΔCTS-LPAT1，并且其中(i)生物体表达FATB硫酯酶，优选拟南芥FATB硫酯酶，经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达AGPAT1，并且其中(i)生物体经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因(FAE1)和脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达AGPAT1，并且其中(i)生物体表达FATB硫酯酶，优选拟南芥FATB硫酯酶，经处理或遗传修饰以破坏脂肪酸去饱和酶2基因(FAD2)，(ii)抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化，及(iii)抑制或阻止生物体内天然内质网溶血磷脂酸酰基转移酶(ER LPAT)的活性。
优选地，通过抑制或阻止磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性来抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化。
优选地，通过抑制或阻止胆碱磷酸转移酶(CPT1)和/或乙醇胺磷酸转移酶(EPT1)的活性来抑制或阻止生物体内二酰基甘油向磷脂酰胆碱的转化。
优选地，生物体经处理或遗传修饰表达FATB硫酯酶基因。
根据本发明的另一个方面，提供一种非人类生物体，优选植物，其中表达叶绿体溶血磷脂酸酰基转移酶(LPAT)，其中所述叶绿体LPAT缺乏功能性叶绿体靶向信号。
如上所述，值得注意的是，当阻止N-端叶绿体靶向信号发挥作用时，发现叶绿体LPAT重新定位到内质网，其在此处将C16：0酯化到甘油骨架上的sn-2位。
优选地，生物体经处理或遗传修饰破坏叶绿体靶向序列。
优选地，叶绿体LPAT经处理或遗传修饰破坏叶绿体靶向序列。
优选地，叶绿体LPAT是缺少至少一部分、优选全部叶绿体靶向序列的修饰叶绿体LPAT，优选是缺少至少一部分、优选全部叶绿体靶向序列的修饰植物叶绿体LPAT。
优选地，叶绿体靶向序列是N-末端叶绿体靶向序列。
优选地，生物体经修饰或处理(例如通过遗传修饰，优选通过叶绿体靶向序列的遗传修饰)破坏叶绿体LPAT叶绿体靶向序列。
优选地，所述遗传修饰包括添加或删除核酸和/或氨基酸残基。
优选地，抑制或阻止生物体中天然内质网(ER)LPAT的活性。
优选地，抑制或阻止生物体中一种或多种天然内质网(ER)LPAT的活性。
优选地，非人类生物体是多倍体。
在本说明书中，“抑制”是指减少但不阻止。
优选地，抑制指的是减少至少约50％，优选至少约60％，优选至少约70％，优选至少约80％，优选至少约85％，优选至少约90％，优选至少约95％，优选至少约98％。
优选地，抑制指的是减少约83％。
优选地，抑制指的是减少约50％至约98％，优选约60％至约95％，优选约70％至约90％，优选约80％至约90％。
优选地，C16：0LPAT经密码子优化以在生物体中表达。
优选地，C16：0LPAT经密码子优化以在植物、真菌、酵母或藻类中表达。最优选地，溶血磷脂酸酰基转移酶经密码子优化以在植物或酵母中表达。
优选地，“修饰的叶绿体溶血磷脂酸酰基转移酶(LPAT)”是指由与SEQ ID NO：1或SEQ ID NO：38，优选SEQ ID NO：1具有至少约80％序列一致性，优选与SEQ ID NO：1或SEQID NO：38，优选SEQ ID NO：1具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“修饰的叶绿体溶血磷脂酸酰基转移酶(LPAT)”是指包含与SEQ ID NO：2具有至少约80％序列一致性，优选与SEQ ID NO：2具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“修饰的叶绿体溶血磷脂酸酰基转移酶(LPAT)”是指由SEQ ID NO：1或SEQID NO：38的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：2的氨基酸序列的蛋白质。
优选地，修饰的叶绿体LPAT由种子特异性启动子控制表达。
优选地，修饰的叶绿体LPAT由种子特异性启动子大豆球蛋白-1启动子(ProGLY)控制表达。
优选地，修饰的叶绿体LPAT是缺乏功能性叶绿体靶向信号的截短叶绿体LPAT。
优选地，生物体在ER LPAT翻译起始位点的非编码区5'中包含破坏性插入。优选地，破坏性插入位于ER LPAT翻译起始位点的约139bp 5'。优选地，破坏性插入是ER LPAT翻译起始位点的139bp 5'。优选地，破坏性插入是T-DNA插入。
优选地，生物体中磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶(PDCT)的活性受到抑制或阻止。
优选地，“AGPAT1”是指包含与SEQ ID NO：40具有至少约80％序列一致性，优选与SEQ ID NO：40具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“AGPAT1”是指由与SEQ ID NO：26、37或39具有至少约80％序列一致性，优选与SEQ ID NO：26、37或39具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“AGPAT1”是指由SEQ ID NO：26、37或39的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：40的氨基酸序列的蛋白质。
优选地，“CreLPAT”是指包含与SEQ ID NO：42具有至少约80％序列一致性，优选与SEQ ID NO：42具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“CreLPAT”是指由与SEQ ID NO：36或SEQ ID NO：41具有至少约80％序列一致性，优选与SEQ ID NO：36或SEQ ID NO：41具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“CreLPAT”是指由SEQ ID NO：36或SEQ ID NO：41的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：42的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT2”是指包含与SEQ ID NO：44具有至少约80％序列一致性，优选与SEQ ID NO：44具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT2”是指由与SEQ ID NO：43具有至少约80％序列一致性，优选与SEQ ID NO：43具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“微拟球藻属LPAT2”是指由SEQ ID NO：43的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：44的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT3”是指包含与SEQ ID NO：46具有至少约80％序列一致性，优选与SEQ ID NO：46具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT3”是指由与SEQ ID NO：45具有至少约80％序列一致性，优选与SEQ ID NO：45具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“微拟球藻属LPAT3”是指由SEQ ID NO：45的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：46的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT4”是指包含与SEQ ID NO：48具有至少约80％序列一致性，优选与SEQ ID NO：48具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“微拟球藻属LPAT4”是指由与SEQ ID NO：47具有至少约80％序列一致性，优选与SEQ ID NO：47具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“微拟球藻属LPAT4”是指由SEQ ID NO：47的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：48的氨基酸序列的蛋白质。
优选地，“集胞藻属LPAT”是指包含与SEQ ID NO：50具有至少约80％序列一致性，优选与SEQ ID NO：50具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“集胞藻属LPAT”是指由与SEQ ID NO：49具有至少约80％序列一致性，优选与SEQ ID NO：49具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“集胞藻属LPAT”是指由SEQ ID NO：49的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：50的氨基酸序列的蛋白质。
优选地，“天然内质网溶血磷脂酸酰基转移酶”是指包含与SEQ ID NO：52、SEQ IDNO：54、SEQ ID NO：56、SEQ ID NO：58、SEQ ID NO：60或SEQ ID NO：62具有至少约80％序列一致性，优选与SEQ ID NO：52、SEQ ID NO：54、SEQ ID NO：56、SEQ ID NO：58、SEQ ID NO：60或SEQ ID NO：62具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“天然内质网溶血磷脂酸酰基转移酶”是指由与SEQ ID NO：51、SEQ IDNO：53、SEQ ID NO：55、SEQ ID NO：57、SEQ ID NO：59或SEQ ID NO：61具有至少约80％序列一致性，优选与SEQ ID NO：51、SEQ ID NO：53、SEQ ID NO：55、SEQ ID NO：57、SEQ ID NO：59或SEQ ID NO：61具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“天然内质网溶血磷脂酸酰基转移酶”是指由SEQ ID NO：51、SEQ ID NO：53、SEQ ID NO：55、SEQ ID NO：57、SEQ ID NO：59或SEQ ID NO：61的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：52、SEQ ID NO：54、SEQ ID NO：56、SEQ ID NO：58、SEQ ID NO：60或SEQ ID NO：62的氨基酸序列的蛋白质。
优选地，“磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶”是指包含与SEQ ID NO：64、SEQ ID NO：66、SEQ ID NO：68、SEQ ID NO：70或SEQ ID NO：72具有至少约80％序列一致性，优选与SEQ ID NO：64、SEQ ID NO：66、SEQ ID NO：68、SEQ ID NO：70或SEQ ID NO：72具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶”是指由与SEQ ID NO：63、SEQID NO：65、SEQ ID NO：67、SEQ ID NO：69或SEQ ID NO：71具有至少约80％序列一致性，优选与SEQ ID NO：63、SEQ ID NO：65、SEQ ID NO：67、SEQ ID NO：69或SEQ ID NO：71具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶”是指由SEQ ID NO：63、SEQ IDNO：65、SEQ ID NO：67、SEQ ID NO：69或SEQ ID NO：71的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：64、SEQ ID NO：66、SEQ ID NO：68、SEQ ID NO：70或SEQ ID NO：72的氨基酸序列的蛋白质。
优选地，“拟南芥FATB硫酯酶”是指包含与SEQ ID NO：73具有至少约80％序列一致性，优选与SEQ ID NO：73具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“拟南芥FATB硫酯酶”是指由与SEQ ID NO：33具有至少约80％序列一致性，优选与SEQ ID NO：33具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“拟南芥FATB硫酯酶”是指由SEQ ID NO：33的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：73的氨基酸序列的蛋白质。
优选地，“FAB1”是指包含与SEQ ID NO：79具有至少约80％序列一致性，优选与SEQID NO：79具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“FAB1”是指由与SEQ ID NO：78具有至少约80％序列一致性，优选与SEQID NO：78具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“FAB1”是指由SEQ ID NO：78的核苷酸序列编码的蛋白质和/或包含SEQID NO：79的氨基酸序列的蛋白质。
优选地，“FAD2”是指包含与SEQ ID NO：81或85具有至少约80％序列一致性，优选与SEQ ID NO：81或85具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“FAD2”是指由与SEQ ID NO：80或84具有至少约80％序列一致性，优选与SEQ ID NO：80或84具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“FAD2”是指由SEQ ID NO：80或84的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：81或85的氨基酸序列的蛋白质。
优选地，“FAE1”是指包含与SEQ ID NO：83或87具有至少约80％序列一致性，优选与SEQ ID NO：83或87具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“FAE1”是指由与SEQ ID NO：82或86具有至少约80％序列一致性，优选与SEQ ID NO：82或86具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“FAE1”是指由SEQ ID NO：83或87的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：82或86的氨基酸序列的蛋白质。
优选地，“胆碱磷酸转移酶(CPT1)”是指包含与SEQ ID NO：89具有至少约80％序列一致性，优选与SEQ ID NO：89具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“胆碱磷酸转移酶(CPT1)”是指由与SEQ ID NO：88具有至少约80％序列一致性，优选与SEQ ID NO：88具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“胆碱磷酸转移酶(CPT1)”是指由SEQ ID NO：88的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：89的氨基酸序列的蛋白质。
优选地，“乙醇胺磷酸转移酶(EPT1)”是指包含与SEQ ID NO：91具有至少约80％序列一致性，优选与SEQ ID NO：91具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的氨基酸序列的蛋白质。
优选地，“乙醇胺磷酸转移酶(EPT1)”是指由与SEQ ID NO：90具有至少约80％序列一致性，优选与SEQ ID NO：90具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的核苷酸序列编码的蛋白质。
优选地，“乙醇胺磷酸转移酶(EPT1)”是指由SEQ ID NO：90的核苷酸序列编码的蛋白质和/或包含SEQ ID NO：91的氨基酸序列的蛋白质。
根据本发明的另一个方面，提供本文所述的非人类生物体的细胞。优选地，细胞是重组细胞。
根据本发明的另一个方面，提供用于产生本文所述植物的种子。
根据本发明的另一个方面，提供从本文所述植物收获的种子、果实或叶子。
根据本发明的另一个方面，提供从本文所述非人类生物体或其细胞产生的甘油三酯。优选地，提供由本文所述植物，优选地，植物的种子、果实或叶子产生的甘油三酯。
优选地，甘油三酯在sn-2位包含约30％以上的C16：0，优选在sn-2位包含约35％以上，优选约40％以上，优选约45％以上，优选约50％以上，优选约55％以上，优选约60％以上，优选约65％以上，最优选约70％以上的C16：0。
优选地，甘油三酯在sn-2位包含约30％至约100％的C16：0，优选在sn-2位包含约35％至约100％，优选约40％至约100％，优选约45％至约100％，优选约50％至约100％，优选约55％至约100％，优选约60％至约100％，优选约65％至约100％，最优选约70％至约100％的C16：0。
优选地，生物体是植物，甘油三酯是从植物的种子、果实和/或叶子获得。
根据本发明的另一个方面，提供从本文所述的生物体，优选本文所述的植物提取甘油三酯的方法，优选其中所述方法包括机械提取和/或溶剂提取。
根据本发明的另一个方面，提供从本文所述的酵母产生甘油三酯的方法，所述方法包括培养所述酵母并从其中提取甘油三酯。
在本发明的另一个方面，提供生产婴儿配方奶粉的方法，包括从本文所述的非人类生物体或其细胞获得甘油三酯，并使用所述甘油三酯生产婴儿配方奶粉。
优选地，生物体是植物，甘油三酯从植物中提取。
优选地，甘油三酯从植物的种子、果实和/或叶子提取。
根据本发明的另一个方面，提供包含如本文所述的和/或由本文所述方法生产的甘油三酯的婴儿配方奶粉。
根据本发明的另一个方面，提供了包含C16：0LPAT的非人类生物体，其中生物体在包含碳源的培养基中培养，其中所述碳源包含(i)一种或多种糖和(ii)一种或多种脂肪酸和/或脂肪酸酯的混合物。
优选地，碳源包含(i)一种或多种糖和(ii)一种或多种油，优选植物油的混合物。
优选地，一种或多种糖包括一种或多种可发酵糖。
优选地，一种或多种糖选自木糖、乳糖、纤维素、葡萄糖、果糖、蔗糖或水解的木质纤维素材料中的一种或多种。
优选地，一种或多种脂肪酸和/或脂肪酸酯包含C16：0。
优选地，一种或多种脂肪酸和/或脂肪酸酯包括脂肪酸和/或脂肪酸酯的混合物，其中至少约30％w/w的脂肪酸和/或脂肪酸酯包含C16：0和/或至少约30％w/w的脂肪酸和/或脂肪酸酯包含C18：1。
优选地，碳源包含棕榈油与葡萄糖和/或甘油的组合。
优选地，生物体经处理或遗传修饰表达C16：0LPAT。
优选地，C16：0LPAT是异源C16：0LPAT。
正如将理解的那样，甘油三酯作为其中的成分构成婴儿配方奶粉的一部分。
优选地，婴儿配方奶粉包含一种或多种其它成分。优选地，所述一种或多种其它成分包括水、乳糖、乳化剂、益生元、益生菌、维生素和/或矿物质中的一种或多种。
在本说明书中，采用使说明书编写清晰简明的方式对实施例进行了描述，但其目的及将理解的是，这些实施方案可以以各种方式组合或分开，但并不背离本发明。例如，应当理解的是，本文描述的所有优选特征都适用于本文描述的本发明的所有方面。
附图说明
现在将参照附图描述本发明的示例性实施例，其中
图1A示出了拟南芥中甘油脂从头(de novo)生物合成的胞质和叶绿体途径。三个修饰使棕榈酰(C16：0)基团(白柱)能够在种子发育过程中结合到TAG的sn-2(或β)位。(1)将LPAT1重新靶向到ER，(2)敲低LPAT2和(3)敲除PDCT。C18：x，长链单-或多不饱和脂肪酰基(黑柱)；CoA，辅酶A；ACP，酰基载体蛋白；G3P，3-磷酸甘油；1-LPA，sn-1溶血磷脂酸；PA，磷脂酸；DAG，二酰基甘油；TAG，甘油三酯；PC，磷脂酰胆碱；1-LPC，sn-1溶血磷脂酰胆碱；FFA，游离脂肪酸；LPAT，1-LPA酰基转移酶；PDCT，PC：DAG胆碱磷酸转移酶；
图1B示出了ΔCTS-LPAT1的DNA序列(SEQ ID NO：1)和氨基酸序列(SEQ ID NO：2)；
图2示出了叶绿体LPAT1可以重新靶向胞质甘油脂生物合成途径，以将C16：0结合到TAG的sn-2位。(A)瞬时表达RFP-ΔCTS-LPAT1和m-GFP5-ER标记的本氏烟表皮细胞的激光扫描共聚焦显微图像。比例尺＝20μm。(B)拟南芥中种子特异性ΔCTS-LPAT1表达对在TAG的sn-2位酯化的C16：0百分数的影响，与sn-1+3位对比。WT＝野生型；L30、L6和L11＝三个独立的纯合ProGLY：ΔCTS-LPAT1系。数值是每种基因型三株植物的不同种子批次的测量平均值±SE(n＝3)。a、b和c表示与WT显著不同(P<0.05)的数值(ANOVA+Tukey HSD检验)；
图3示出了ER-驻留LPAT2的破坏增加了C16：0结合到TAG的sn-2位。(A)LPAT2位点图，示出了突变等位基因中T-DNA插入的位置。lpat2突变体背景对(B)在TAG的sn-2位酯化的C16：0百分含量的影响(与sn-1+3位比较)和(C)表达ΔCTS-LPAT1的种子中LPAT2转录本丰度的影响。WT＝野生型；L11＝纯合ProGLY：ΔCTS-LPAT1系。数值是每种基因型三株植物(n＝3)的不同批次干种子(B中)和正在发育的长角果(C中)的测量平均值±SE。LPAT2表达被归一化为三个参考基因的几何平均值，并相对于WT表达。a、b和c表示与L11显著不同(P<0.05)的数值(ANOVA+Tukey HSD检验)；
图4示出了旁路通过PC的通量增加了C16：0结合到TAG的sn-2位。(A)pdct突变体背景对ProGLY：ΔCTS-LPAT1和ProGLY：ΔCTS-LPAT1 lpat2-3种子中在TAG的sn-2位酯化的C16：0百分含量的影响。WT＝野生型；L11＝纯合ProGLY：ΔCTS-LPAT1系。(B)种子重量和(C)WT和ProGLY：ΔCTS-LPAT1 lpat2-3pdct(全部)的含油量。数值是每种基因型3到6株植物(A中)及5株植物(B和C中)(n＝3-6)的不同种子批次的测量平均值±SE。a和b分别表示与L11和pdct显著不同(P<0.05)的数值(ANOVA+Tukey HSD检验)，c表示与WT显著不同的数值(双尾学生t检验(two-tailed Student’s t test))；
图5示出了遗传修饰对种子活力的影响。(A)种子发芽率，(B)第4天子叶展开的百分数和(C)第7天长出真叶的百分数。(D)子叶展开和长出真叶的幼苗的代表性图。(E)第0天和第4天种子/幼苗的TAG含量。WT＝野生型；全部(All)＝ProGLY：ΔCTS-LPAT1lpat2-3pdct。数值是每种基因型三株植物(n＝3)不同批次种子的测量平均值±SE。在D中，比例尺＝2mm。a和b表示与WT显著不同(P<0.05)的数值(双尾学生t检验)；
图6示出了遗传修饰对种子脂质组成的影响。(A)三棕榈酸甘油酯含量和(B)PC组成。WT＝野生型；全部(All)＝ProGLY：ΔCTS-LPAT1 lpat2-3pdct。数值是每种基因型五株植物的不同种子批次的测量平均值±SE(n＝5)。a表示与WT显著不同(P<0.05)的数值(双尾学生t检验)；
图7示出了野生型(WT)和ProGLY：ΔCTS-LPAT1 lpat2-3pdct(全部)植物在开始开花时的图像；
图8简单阐明了在拟南芥种子中产生OPO的策略。采用亚效fab1-1和无效fae1和fad2突变等位基因组合在种子中产生高含量的16：0和18：1。然后使用具有16：0-CoA偏好的ER定位LPAT与亚效lpat2-3和无效pdct突变等位基因的表达，使16：0在1-LPA的sn-2位优先酯化和使产物导入到TG内。16：0，棕榈酸；18：1，油酸；CoA，辅酶A；G3P，3-磷酸甘油；1-LPA，sn-1溶血磷脂酸；PA，磷脂酸；DG，二酰基甘油；TAG，甘油三酯；PC，磷脂酰胆碱；1-LPC，sn-1溶血磷脂酰胆碱；FA，脂肪酸；
图9示出了HPHO种子的TAG的脂肪酸组成。从WT、fab1-1 fae1、fad2 fae1和fab1-1fae1 fad2种子中分离的TAG的总脂肪酸组成。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；
图10示出了WT种子和表达ΔCTS-LPAT1的HPHO种子的TAG中16：0的含量。示出了从WT、ProGLY：ΔCTS-LPAT1、fab1-1 fad2 fae1和ProGLY：ΔCTS-LPAT1 fab1-1 fad2 fae1种子分离的TAG中sn-2位酯化的16：0的百分含量(A)和16：0占总脂肪酸的百分含量(B)。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；*表示与WT显著(P<0.05)不同的数值，或在括号中标记的情况下，表示数值彼此显著不同(ANOVA+Tukey HSD检验)；
图11示出了WT种子和表达AGPAT1的HPHO种子的TAG中16：0的含量。瞬时表达RFP-AGPAT1和m-GFP5-ER标记的本氏烟表皮细胞的激光扫描共聚焦显微图像(A)。比例尺＝20μm。示出了从WT、ProGLY：AGPAT1、fab1-1 fad2 fae1和ProGLY：AGPAT1 fab1-1 fad2 fae1种子分离的TAG中sn-2位酯化的16：0的百分含量(B)和16：0占总脂肪酸的百分含量(C)。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；*表示与WT显著(P<0.05)不同的数值，或在括号中标记的情况下，表示数值彼此显著不同(ANOVA+Tukey HSD检验)；
图12示出了来自表达ΔCTS-LPAT1或AGPAT1的HPHO lpat2-3 pdct种子的TAG中16：0的含量。从WT、ProGLY：ΔCTS-LPAT1 fab1-1 fae1 fad2、lpat2-3 pdct和ProGLY：AGPAT1 fab1-1 fae1 fad2 lpat2-3 pdct种子分离的TAG中sn-2位酯化的16：0的百分含量(A)和16：0占总脂肪酸的百分含量(B)。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；*表示与WT显著(P<0.05)不同的数值，或在括号中标记的情况下，表示数值彼此显著不同(ANOVA+Tukey HSD检验)；
图13示出了表达ΔCTS-LPAT1或AGPAT1的HPHO lpat2-3 pdct种子的油含量。WT、fab1-1 fae1 fad2、ProGLY：ΔCTS-LPAT1 fab1-1 fae1 fad2、lpat2-3 pdct和ProGLY：AGPAT1 fab1-1 fae1 fad2 lpat2-3 pdct种子的种子重量(A)和百分含油量(B)。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；*表示与WT显著不同的数值(P<0.05)(ANOVA+Tukey HSD检验)；和
图14示出了表达ΔCTS-LPAT1或AGPAT1的HPHO lpat2-3 pdct种子的活力。WT、fab1-1 fae1 fad2、ProGLY：ΔCTS-LPAT1 fab1-1 fae1 fad2、lpat2-3 pdct和ProGLY：AGPAT1 fab1-1 fae1 fad2 lpat2-3 pdct种子的发芽率(A)和子叶在第4天完全展开的百分数(B)。数值是每种基因型三株植物(n＝3)的种子的测量平均值±SE；*表示与WT显著不同(P<0.05)的数值(ANOVA+Tukey HSD检验)。
具体实施方式
本发明涉及用于产生甘油三酯的非人类生物体，其中所述甘油三酯模拟人类甘油三酯的脂肪酸立体异构结构。特别是，本发明提供产生甘油三酯的非人类生物体，所述甘油三酯在甘油骨架sn-2位的C16：0百分含量与人类甘油三酯相似。
本文所述的本发明的目的是探索植物油的立体异构结构是否可以通过迭代代谢工程改变，从而使其模拟HMF。据我们所知，没有任何陆生植物(胚生植物)在sn-2位(与sn-1/3位比较)产生富含C16：0的TAG，并且在几乎所有情况下，C16：0基本上都被排除在sn-2位之外(4，5，8)。即使在总共含有～48％C16：0的棕榈油中，也只有9％C16：0占据sn-2位。本文描述了一种在拟南芥油籽模型中修饰TAG生物合成的方法，所述方法导致酰基的立体异构重新分布，从而使sn-2位C16：0的含量增加20倍以上，达到总含量的70％以上；这种富集含量与HMF相当。设想将这项技术应用于油料作物，因为其不需要基于酶的催化，因此将为婴儿配方奶粉提供更便宜和因此更广泛可得的HMFS来源。
在本说明书中，术语“约”是指正负20％，更优选正负10％，甚至更优选正负5％，最优选正负2％。
在本说明书中，术语“对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶”是指优先于其他脂肪酰基-CoA底物，优选优先于长链不饱和脂肪酰基-辅酶A，在甘油骨架的sn-2位酯化C16：0的溶血磷脂酸酰基转移酶。换句话说，“对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶”在使用C16：0-辅酶A时比使用其他脂肪酰基-CoA，优选比长链不饱和脂肪酰基-CoA具有更大的活性。
优选地，术语“对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶”是指优先于使用酰基链长超过16个碳原子且含有一个或多个双键的脂肪酰基-CoA底物，在sn-1溶血磷脂酸的甘油骨架上的sn-2位酯化C16：0的溶血磷脂酸酰基转移酶。
在本说明书中，术语“对C16：0-辅酶A不具有特异性的溶血磷脂酸酰基转移酶”是指不优先于其他脂肪酰基-CoA底物，优选不优先于长链不饱和脂肪酰基-辅酶A，在甘油骨架的sn-2位酯化C16：0的溶血磷脂酸酰基转移酶。换句话说，“对C16：0-辅酶A不具有特异性的溶血磷脂酸酰基转移酶”在使用C16：0-辅酶A时比使用其他脂肪酰基-CoA，优选比长链不饱和脂肪酰基-CoA具有更低的活性。
优选地，术语“对C16：0-辅酶A不具有特异性的溶血磷脂酸酰基转移酶”是指不优先于使用酰基链长超过16个碳原子且含有一个或多个双键的脂肪酰基-CoA底物，在sn-1溶血磷脂酸的甘油骨架上的sn-2位酯化C16：0的溶血磷脂酸酰基转移酶。
在本说明书中，正如本领域所了解的那样，“一致性”是如通过比较序列确定的两个或更多个多肽序列或两个或更多个多核苷酸序列之间的关系。在本领域中，“一致性”还指的是多肽或多核苷酸序列之间的序列相关程度，视情况而定，由此类序列的字符串之间的匹配确定。一致性百分数可以很容易地通过已知方法计算，包括但不限于ComputationalMolecular Biology,Lesk,A.M.,ed.,Oxford University Press,New York,1988；Biocomputing:Informatics and Genome Projects,Smith,D.W.,ed.,Academic Press,New York,1993；Computer Analysis of Sequence Data,Part I,Griffin,A.M.,andGriffin,H.G.,eds.,Humana Press,New Jersey,1994；Sequence Analysis in MolecularBiology,von Heinje,G.,Academic Press,1987；和Sequence Analysis Primer,Gribskov,M.and Devereux,J.,eds.,M Stockton Press,New York,1991；和Carillo,H.,and Lipman,D.,SLAM J.Applied Math.,48:1073(1988)中描述的方法，通过引用，将所有这些方法整体并入本发明中。确定一致性的优选方法设计用于给出测试序列之间的最大匹配度。确定一致性的方法被编入公开可用的计算机程序中。确定两个序列之间一致性百分数的优选计算机程序方法包括但不限于GCG程序包(Devereux,J.,et al.,Nucleic AcidsResearch 12(1):387(1984)，通过引用，将其整体并入本发明中)、BLASTP、BLASTN和FASTA((Atschul,S.F.et al.,J.Molec.Biol.215:403-410(1990)，通过引用，将它们整体并入本发明中)。BLAST X程序可从NCBI和其他来源公开获得(BLAST Manual,Altschul,S.,et al,NCBI NLM NIH Bethesda,Md.20894；Altschul,S.,et al.,J.Mol.Biol.215:403-410(1990)，通过引用，将它们整体并入本发明中)。作为说明，多核苷酸具有与参考核苷酸序列“SEQ ID NO：A”至少例如95％“一致性”的核苷酸序列，指的是除了在“SEQ ID NO：A”参考核苷酸序列的每100个核苷酸中可包含至多5个点突变之外，该多核苷酸的核苷酸序列与参考核苷酸序列相同。换句话说，为了获得具有与参考核苷酸序列至少95％一致性的核苷酸序列的多核苷酸，参考序列中不超过5％的核苷酸可以删除或用另一种核苷酸替代，或可以将数量不超过参考序列中总核苷酸5％的多个核苷酸插入到参考序列中。参考序列的这些突变可以发生在参考核苷酸序列的5'或3'末端位置或这些末端位置之间的任何位置，单独散布在参考序列的各个核苷酸之间或散布在参考序列的一个或多个连续组中。类似地，多肽具有与参考氨基酸序列“SEQ ID NO：B”至少例如95％“一致性”的氨基酸序列，指的是除了在“SEQ ID NO：B”参考氨基酸序列的每100个氨基酸中可包含不超过5个氨基酸变动之外，该多肽的氨基酸序列与参考序列相同。换句话说，为了获得具有与参考氨基酸序列至少95％一致性的氨基酸序列的多肽，参考序列中不超过5％的氨基酸残基可以删除或用另一种氨基酸替代，或可以将数量不超过参考序列中总氨基酸残基5％的多个氨基酸插入到参考序列中。参考序列的这些变动可以发生在参考氨基酸序列的氨基或羧基末端位置或这些末端位置之间的任何位置，单独散布在参考序列的各个残基之间或散布在参考序列的一个或多个连续组中。
关于本文所述的序列一致性，应当理解的是，本发明涉及的许多非人类生物体是多倍体。这意味着它们可以拥有序列一致性差异很小的每个基因的多个拷贝。因此，应当理解的是，本文所述的特定基因或蛋白质还包括该基因和来自同一生物体的相关编码蛋白质的一个或多个其它拷贝。
优选非人类生物体是多倍体，本文提到的特定序列包括与所述序列具有至少约80％序列一致性的序列，优选与所述序列具有至少约85％序列一致性，优选至少约90％序列一致性，优选至少约95％序列一致性，优选至少约98％序列一致性，优选至少约99％序列一致性的序列。
应当理解的是，“一个或多个”包括“多个”。
在本说明书中，“遗传修饰”是指生物体的DNA已被修饰的生物体。例如，这可以通过从另一个生物体引入DNA或通过修饰现有的DNA来实现。
实施例1
结果与讨论
LPAT1可以通过删除其叶绿体靶向信号重新定向到ER
在植物细胞中，甘油三酯(TAG)由位于内质网(ER)上的胞质甘油脂生物合成途径形成，负责sn-2位酰化的酶是溶血磷脂酸酰基转移酶(LPAT)(图1A)。LPAT的ER驻留同种型会排斥作为底物的C16：0-辅酶A(CoA)，这可能是C16：0被排除在sn-2位之外的原因。为了克服这种限制，我们决定表达一种对C16：0-CoA具有特异性的LPAT(图1A)。已经描述了来自蓝细菌、哺乳动物和藻类的几种候选转基因。然而，植物已经拥有一个驻留在叶绿体中的具有适当选择性的LPAT(图1A)。该LPAT使用C16：0-酰基载体蛋白(ACP)底物，但在体外也接受C16：0-CoA。因此，我们决定测试叶绿体LPAT是否可以重新定位到ER(图1A)。叶绿体LPAT是一种完整的膜蛋白，由核编码，并包含N-端叶绿体靶向信号(CTS)。利用本氏烟叶中的瞬时表达，我们发现，当从欧洲油菜LPAT1中删除101个含有CTS的氨基酸残基(图1B)并采用红色荧光蛋白(RFP)标记取代时，RFP-ΔCTS-LPAT1融合蛋白定位到ER(图2A)。
ΔCTS-LPAT1表达促进C16：0结合到TAG的sn-2位
据悉，缺乏CTS的截短型LPAT1在大肠杆菌中表达时具有活性。为了确定ΔCTS-LPAT1是否在植物中发挥作用，并使C16：0能够结合到TAG的sn-2位，我们在拟南芥油籽模型中在种子特异性大豆球蛋白-1启动子(ProGLY)的控制下表达了这种截短的蛋白质。我们使用DsRed荧光标记系统选择了40多个初级转化体(T1)，并分析了T2种子批次的总脂肪酰基组成。我们发现几个系表现出总C16：0含量增加，这表明转基因促进C16：0结合到TAG内(表1)。
表1.各野生型(WT)和分离T2 ProGLY：ΔCTS-LPAT1系(L1-42)各种子批次的总脂
肪酸组成。星号表示供进一步分析的单拷贝系。
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我们选择了C16：0含量高的三个独立的单拷贝T2系(L30、L6和L11)，获得了纯合T3种子。当我们从这些纯合种子批次纯化TAG，并使用脂肪酶消化测定其立体化学时，我们发现，sn-2位(相对于sn-1+3位)的C16：0百分含量增加了16倍以上，从野生型仅约2％至三个独立的ProGLY：ΔCTS-LPAT1系增加到～32％至～39％(图2B；表2)。
表2.野生型(WT)种子和纯合ProGLY：ΔCTS-LPAT1种子中TAG的总脂肪酰基和sn-2
脂肪酰基组成。
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因此，ΔCTS-LPAT1表达足以允许将C16：0结合到TAG的sn-2位，此位可以结合低比例的C16：0，但与sn-1/3位相比，无法在此位实现正富集(图1A)。
LPAT2的破坏增强C16：0结合到TAG的sn-2位
异源和天然酰基转移酶之间的竞争可能是限制特定脂肪酰基结合到TAG的一个因素。因此，我们研究了是否可以通过破坏天然ER驻留LPAT(被认为是拟南芥中的LPAT2)的功能来增强C16：0的ΔCTS-LPAT1依赖性结合到TAG的sn-2位(图1A)。lpat2-1无效突变体是胚胎致死的。然而，在必需基因的非编码区的T-DNA插入可用于产生可行的亚效等位基因。因此，我们分离了两个T-DNA突变体(lpat2-2和lpat2-3)，这两个突变体分别插入302和139bpLPAT2翻译起始位点的5'(图3A)。然后，我们将ProGLY：ΔCTS-LPAT1 L11杂交到每个新的lpat2等位基因中，并回收纯合种子批次。当我们从这些种子批次中纯化TAG并进行位置分析时，我们发现sn-2位的C16：0百分含量从亲本ProGLY：ΔCTS-LPAT1系中的～33％增加到lpat2-3背景中的～51％，而lpat2-2背景中的效果不显著(P>0.05)(图3B；表3)。
表3.野生型(WT)种子和含lpat2突变等位基因的纯合ProGLY：ΔCTS-LPAT1种子中
TAG的总脂肪酰基和sn-2脂肪酰基组成。
qRT-PCR分析表明，LPAT2表达在生长中的lpat2-3长角果中减少～83％，但在lpat2-2中仅减少～24％。(图3B)。这些数据支持这一假设：LPAT2有助于拟南芥种子中的TAG生物合成，并且它与ΔCTS-LPAT1竞争。sn-2位C16：0的富集水平似乎也对LPAT2抑制强度有反应，因此，实现大于～83％的LPAT表达减少可能会导致更强的富集。
PDCT的破坏也增强C16：0结合到TAG的sn-2位
在拟南芥种子的发育过程中，TAG中>90％的甘油骨架来源于膜脂磷脂酰胆碱(PC)，这是由于二酰基甘油(DAG)-PC之间的快速相互转化导致的，这种相互转化主要由植物特异性头部基团交换酶PC：DAG胆碱磷酸转移酶催化(PDCT)(图1A)。尽管LPAT负责甘油脂在sn-2位的初始酰化，但一旦这些酰基在PC中，它们可能会被被酰基编辑活动删除和取代(图1A)。为了确定旁路通过PC的甘油脂通量(图1A)是否可能增加C16：0的ΔCTS-LPAT1依赖性结合到TAG的sn-2位，我们将ProGLY：ΔCTS-LPAT1 L11杂交到pdct(减少油酸去饱和1)突变体内。当我们从ProGLY：ΔCTS-LPAT1 pdct种子批次中纯化TAG并进行位置分析时，我们发现，sn-2位处C16：0的百分含量从亲本ProGLY：ΔCTS-LPAT1系的～30％增加到pdct背景中的～56％(图4A；表4)。
表4.野生型(WT)种子和含rod1和lpat2-3突变等位基因的纯合ProGLY：ΔCTS-
LPAT1种子中TAG的总脂肪酰基和sn-2脂肪酰基组成。
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这些数据表明，新制造的DAG更直接地进入TAG(图1A)有利于C16：0结合和/或保留在sn-2位。在WT种子中，可以设想进入PC的sn-2位的C16：0可能通过溶血磷脂酰胆碱酰基转移酶(LPCAT)或磷脂酶A2的作用从中编辑。有趣的是，一些研究小组提供了体外证据，表明拟南芥LPCAT1和LPCAT2的反向活性可以选择性地从PC中删除某些脂肪酰基，但未对C16：0进行试验。
LPAT2和PDCT的破坏对C16：0在sn-2位的结合具有加性效应
为了确定减少LPAT竞争和旁路通过PC的通量相组合是否会对C16：0的ΔCTS-LPAT1依赖性结合到TAG的sn-2位产生加性效应(图1A)，我们将ProGLY：ΔCTS-LPAT1lpat2-3与ProGLY：ΔCTS-LPAT1 pdct进行杂交。当我们从纯合种子批次中纯化TAG并进行位置分析时，我们发现，sn-2位C16：0的百分含量从ProGLY：ΔCTS-LPAT1 pdct的～56％增加到ProGLY：ΔCTS-LPAT1 lpat2-3 pdct中的～71％(图4A；表4)。因此，对拟南芥中TAG生物合成途径的三种修饰(即ΔCTS-LPAT1表达，加上LPAT2和PDCT抑制)的组合足以复制HMF中sn-2位的C16：0富集水平(相对于sn-1+3位)。使用高分辨率/精确质量(HR/AM)脂质组学分析ProGLY：ΔCTS-LPAT1 lpat2-3pdct(全部)种子中的TAG组成，也证实了在sn-2位存在C16：0基团，因为与WT相比，三棕榈酸甘油酯的富集高27-倍(图6A)。相比之下，在ProGLY：ΔCTS-LPAT1 lpat2-3pdct种子中没有检测到二棕榈酰PC，并且含有一个C16：0基团的PC分子种类没有增加(图6B)。这些数据表明，PC中饱和及不饱和脂肪酰基的不对称分布在ProGLY：ΔCTS-LPAT1 lpat2-3pdct种子中得到保持，这对于防止膜在生理温度下呈现凝胶相可能很重要。
C16：0的重新分布降低了种子含油量，但未降低发芽率或成苗率
许多研究表明，修饰脂肪酰基组成会减少油籽中TAG的积累，在某些情况下还会损害种子发芽和成苗。我们在这项研究中的主要目标并不是改变脂肪酰基组成本身，而是改变TAG的立体异构结构。为了考察C16：0在TAG的sn-2位富集的生理影响，我们比较了在标准实验室条件下一起生长的野生型和ProGLY：ΔCTS-LPAT1 lpat2-3pdct植物种子批次。我们发现，这两种基因型的种子重量并没有显著差异(P>0.05)(图4B)。但是，当以种子重量百分数表示时，ProGLY：ΔCTS-LPAT1 lpat2-3pdct种子的脂肪酸含量显著(P<0.05)低于野生型(图4C)。这些数据表明，导致C16：0结合到sn-2位的这些修饰降低了TAG生物合成通量。这一发现与之前使用基因工程或突变育种方法改变种子中TAG组成的研究一致。尽管脂肪酸含量降低，但ProGLY：ΔCTS-LPAT1 lpat2-3pdct的种子发芽率(以胚根出现计分)(图5A)、成苗率(以子叶展开计分)(图5B)和真叶生长率(图5C)似乎并未显著(P<0.05)受损。ProGLY：ΔCTS-LPAT1 lpat2-3 pdct种子发芽后TAG的分解也没有受到阻碍(图5D)，这与其中种子经修饰以将不常见的脂肪酰基结合到TAG中的一些研究形成对比。最后，尽管ProGLY：ΔCTS-LPAT1 lpat2-3pdct携带必需基因LPAT2的亚效等位基因，但这似乎不会对莲座期的生长和形态产生不利影响(图7)。
材料和方法
植物材料和生长条件
拟南芥野生型(Col-0)和LPAT2(At3g57650)T-DNA插入系SALK_108026(lpat2-2)和SALK_004681(lpat2-3)(1)从欧洲拟南芥种子中心(英国诺丁汉大学)获得。所述pdct(rod1)突变体先前已进行了描述(2)。种子经过表面消毒，在4℃下分层两天，放在含有pH5.7、1/2强度MS培养基(Sigma-Aldrich)的琼脂板上发芽。将幼苗移植到含有Levington F2堆肥的7cm2盆中，并在设置为16小时光照(22℃)(光照强度为250μmol m-2s-1)/8小时黑暗(16℃)循环的室中生长。在开花开始时将植物单独套袋，并在成熟时收获种子。
克隆和转化
从欧洲油菜(cv.Kumily)幼苗中分离RNA，并使用RNAeasy Plant Mini试剂盒(Qiagen)处理DNAse。使用SuperScript III逆转录酶试剂盒(ThermoFisher Scientific)将RNA逆转录为cDNA。使用引物对P1+P2，采用KOD DNA聚合酶(Merck)通过PCR扩增缺乏叶绿体靶向序列(CTS)的欧洲油菜LPAT1(GenBank：AF111161)(表5)。
表5.研究中使用的引物
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使用QIAquick凝胶回收试剂盒(Qiagen)纯化得到的PCR产物。为了进行定位研究，在pK7WGR2载体(弗拉姆斯生物技术研究所)中的红色荧光蛋白(RFP)后面，克隆ΔCTS-LPAT1。将PCR产物克隆到pENTRTM/D-TOPOTM载体(赛默飞世尔科技(Thermo FisherScientific))中，测序(图1B)，并使用GatewayTM LR ClonaseTM II Enzyme mix(赛默飞世尔科技)重组到pK7WGR2载体中。ΔCTS-LPAT1被克隆到位于大豆球蛋白-1(GLY)启动子和种子特异性表达终止子之间的pBinGlyRed3载体中(3)。ΔCTS-LPAT1使用KOD DNA聚合酶和引物对P3+P4从pENTR-D-TOPO载体进行PCR扩增。PCR产物经凝胶纯化并用EcoRI和XbaI消化。pBinGlyRed3载体也用EcoRI和XbaI消化，碱性磷酸酶处理(普洛麦格(Promega))，凝胶纯化，并使用T4 DNA连接酶(NEB)将ΔCTS-LPAT1连接到该载体中。使用热激将载体转化为根癌农杆菌菌株GV3101，然后使用浸花法(4)进行拟南芥转化。使用DsRed滤光片在LeicaM205 FA显微镜下鉴定表达可选择标记的T1种子。
本氏烟中的瞬时表达和成像
正如Wood et al.,(5)所述，使用含有Pro35S：RFP-ΔCTS-LPAT1、Pro35S：m-GFP5-ER或Pro35S：p19的载体转化的根癌农杆菌培养物在本氏烟叶中进行瞬时表达。将培养物手工渗透到叶子中，并将接种的植物放置48小时。然后在Apochromat 63x/1.20W Korr M27物镜下，将本氏烟叶安装在水中置于Zeiss LSM 780激光扫描共聚焦显微镜上。GFP在488nm的波长处被激发，RFP在561nm处被激发。检测采用具有473-551nm发射带的滤光片。
基因分型
使用DNeasy Plant Mini试剂盒(Qiagen)分离基因组DNA。使用普洛麦格PCRMaster Mix(普洛麦格)通过PCR(1)鉴定纯合T-DNA插入突变体。以下引物对用于基因分型lpat2-2(WT：P5+P6或T-DNA：P5+P7)和lpat2-3(WT：P8+P9或T-DNA：P8+P7)。PDCT通过对使用跨越点突变位点的引物对P10+P11扩增的PCR产物进行测序来进行基因分型(2)。ProGLY：ΔCTS-LPAT1使用跨越ProGLY和ΔCTS-LPAT1的引物对P12+P13通过PCR进行基因分型。
脂质分析
从种子和幼苗中提取总脂质，并如前所述纯化TAG(6)。除了2-单酰基甘油使用己烷：乙醚：乙酸(35：70：1.5，v/v/v)通过薄层色谱法(硅胶60,20 x 20cm；Sigma-Aldrich/Merck)(8)分离之外，TAG立体化学分析按照前面描述的方法(7)通过脂肪酶消化进行。如前所述(9)，使用配备DB-23柱(30mx 0.25mm i.d.x 0.25μm)的7890A GC系统(安捷伦科技公司)，通过连接火焰离子化检测的气相色谱(GC)，对整个种子和纯化脂质中存在的脂肪酰基进行反式甲基化和定量。
使用Vanquish-Q Exactive Plus UPLC-MS/MS系统(赛默飞世尔科技)，通过高分辨率/精确质量(HR/AM)脂质组学(10-12)分析TAG和PC的分子种类组成。工作流程包括使用3μg/μl纯化且在氯仿：甲醇(1：1，v/v)中以1：100稀释的总脂质。添加三棕榈酸甘油酯内标(0.857μM)，并将20μl内标注入到UPLC内。脂质使用Accucore C18(2.1x 150mm,2.6mm)色谱柱(赛默飞世尔科技)温度35℃，自动进样器托盘温度10℃，流速400μl min-1的条件分离。流动相：A＝10mM甲酸铵的50％乙腈溶液+0.1％甲酸，B＝2mM甲酸铵的乙腈：丙-2-醇：水(10：88：2v/v)溶液+0.02％甲酸。洗脱梯度从开始时的35％B运行28分钟到100％运行24分钟。Thermo Q Exactive HESI II探针条件，扫描板使用中探针位置在C。调整条件适用于单独的正运行和负运行，以单一极性运行样品导致更多识别。在正离子和负离子模式下开展140K分辨率的LC/MS及与数据无关的HCD MS2实验(35K分辨率)。全扫描@140000分辨率m/z150-1200，含量最丰富的前15名MS/MS@35,000分辨率，使用条件：隔离窗口1m/z、最大积分时间75ms，动态排除窗口8秒。阶梯式碰撞能量是25、30、40eV，负离子模式将25替换为30eV。鞘气设置为60，辅助气体20，吹扫气体1，喷雾电压在正离子模式下为3.2KV，在负离子模式下进行小幅调整，毛细管温度为320，辅助气体加热器设置为370℃。脂质表征采用LipidSearch 4.2实验工作流程(赛默飞世尔科技)，可能的脂质种类与正离子或负离子加合物分开鉴定。通过组合正离子和负离子注释并将它们合并为单个脂质注释，将每个生物重复的数据在色谱时间窗口内对齐。
qRT-PCR分析
如Mendes et al.,(13)所述，从生长的长角果中分离DNA酶处理的总RNA。使用来自Invitrogen有限公司(英国佩斯利)的SuperScriptTM II RNase H-逆转录酶进行单链cDNA的合成。除了将LPAT2和ΔCTS-LPAT1表达标准化为三个参考基因(UBQ5、EF-1α和ACT8)的几何平均值之外，如前所述(13)开展定量(q)-PCR。选择这些参考基因的原因是由于其在种子发育过程中稳定表达(14)。引物对P14+P15、P16+P17、P18+P19、P20+P21和P22+P23分别用于LPAT2、ΔCTS-LPAT1、UBQ5、EF-1α和ACT8。
种子萌发和成苗分析
将每株植物的大约50粒种子播种在1/2 MS琼脂板上，在4℃下分层两天，然后转移到设置为20℃，16h光照/8h黑暗，PPFD＝150μmol m-2s-1的生长室中。一周内每天在解剖立体显微镜下目测对发芽(胚根出现)、子叶展开和真叶展开进行评分。种子和幼苗还在分层后第零天和第四天收集用于脂质分析。
统计分析
所有实验均使用三到六个生物重复进行，数据以平均值±平均值标准误差(SE)表示。对于统计分析，我们使用单向方差分析(ANOVA)和事后Tukey HSD(真实显著性差异)检验，或使用双尾学生t-检验。
结论
如本文所述，可以对植物中的TAG生物合成途径进行工程改造，从而改变种子储油的立体异构结构以模拟HMF的立体异构结构，其中>70％的C16：0集中在甘油骨架上的中间(sn-2或β)位置。越来越多的证据表明这种结构对婴儿营养有益，但尚未发现其天然存在于植物脂肪中，在天然植物脂肪中，C16：0实际上被排除在sn-2位之外。许多婴儿配方奶粉含有通过酶催化重组植物脂肪制备的HMFS，但这种HMFS的生产成本相对较高；制造在sn-2位具有>70％C16：0的真正模拟物时尤其如此。将我们的技术从模式物种拟南芥向油料作物转化，消除了酶催化需求，可能会为婴儿配方奶粉提供更便宜和更可持续的HMFS来源。据估计，目前婴儿配方奶粉市场每年使用约450,000公吨植物性脂肪，其中约38,000公吨是HMFS。几种油料作物被认为是HMFS生产的可能主体，值得注意的是，已经开发出具有适当脂肪酰基组成的常规向日葵和转基因油菜品种。据临床试验报告，sn-2位C16：0仅仅达到43％即具有益处，而且产品调查发现，这种富集水平在添加了HMFS的婴儿配方奶粉中很常见，因此，我们在本文实现的即使sn-2位C16：0富集程度不太高的油料作物仍然可能是可取的。
进一步的研究如下文详细描述。
实施例2–fab1-1fae1背景
我们将实施例1中描述的ProGLY：ΔCTS-LPAT1 lpat2-3 pdct系杂交到fab1-1fae1突变体(15)内，得到ProGLY：ΔCTS-LPAT1 lpat2-3 pdct fab1-1 fae1。由于编码叶绿体3-酮基-酰基-ACP合酶II(KASII)的FAB1基因的亚效等位基因，这种拟南芥背景在其种子TAG中具有升高的C16：0含量，并且还由于无效脂肪酸延长酶1(FAE1)等位基因而降低了极长链脂肪酸的含量(Nguyen et al.,2010)。当我们进行位置分析时，我们发现sn-2位C16：0的百分含量在fab1-1 fae1中是～2％，在ProGLY：ΔCTS-LPAT1 lpat2-3 pdct fab1-1fae1中是～60％(表6)。数据表明，TAG中sn-2位的C16：0富集可以在总C16：0含量高于WT种子的种子中产生。
表6.拟南芥fab1-1 fae种子和ProGLY：ΔCTS-LPAT1 lpat2-3 pdct fab1-1 fae1
种子TAG中C16：0的总含量和sn-2位C16：0的含量。数值是每种基因型两株植物的不同种子
批次的测量值。
实施例3-人LPAT AGPAT1
我们按照实施例1中描述的程序，在大豆球蛋白启动子控制下，在野生型(WT)拟南芥Col-0种子和fab1-1 fae1种子中表达了植物密码子优化型人LPAT AGPAT1。当我们进行位置分析时，我们发现，sn-2位C16：0的百分含量在WT中高达～73％，在fab1-1 fae1背景中高达～55％(表7)。数据表明，TAG中sn-2位的C16：0富集可以通过AGPAT1的表达在种子中产生。
表7.拟南芥WT种子、fab1-1 fae1种子、ProGLY：AGPAT1种子和ProGLY：AGPAT1 L3
fab1-1 fae1种子TAG中C16：0的总含量和sn-2位C16：0的含量。数值是每种基因型三株植物
的不同种子批次的测量平均值±SE。
密码子优化的AGPAT1 DNA序列
实施例4-亚麻荠
我们在亚麻荠中表达了ΔCTS-LPAT1，同时抑制内源性ER LPAT。我们分别在种子特异性油质蛋白和油菜籽蛋白启动子控制下，在包含ΔCTS-LPAT1和LPAT2样基因特异性人工microRNA(amiRNA)的pBinGlyRed3骨架(3)中构建了多基因T-DNA载体。选择三个amiRNA序列靶向亚麻荠LPAT2-样基因(5’-TAAAGCGAGTTCCCTCGACAG-3’(SEQ ID NO：27)、5’-TTGTGCCCAGTGTACGGACTT-3’(SEQ ID NO：28)和5’-TCAAAGGCACGATGATACCTG-3’(SEQ IDNO：29))，并用于取代拟南芥MIR319a前体中的茎环(16)。采用根癌农杆菌将该构建体转化到亚麻荠cv Suneson(WT)中(17)。获得了多个独立系的纯合T3种子批次，并如实施例1中所述开展脂质分析。当我们进行位置分析时，我们发现sn-2位C16：0的百分含量在WT种子中是～1％和在ProOLE：ΔCTS-LPAT1ProNAP：LPAT2amiRNA种子中是～70％(表8)。数据表明，TAG中sn-2位的C16：0富集可以在亚麻荠种子中产生。
表8.亚麻荠WT种子和ProOLE：ΔCTS-LPAT1 ProNAP：LPAT2amiRNA种子TAG中C16：0
的总含量和sn-2位C16：0的含量。数值是每个基因型单个种子批次的测量值。
实施例5-欧洲油菜
我们在欧洲油菜种子中表达ΔCTS-LPAT1或AGPAT1，同时抑制内源性ER LPAT，还表达拟南芥FATB，以增加C16：0总含量。我们分别在种子特异性基因启动子油质蛋白、油菜籽蛋白或大豆球蛋白控制下，在包含FATB、LPAT2-样基因特异性amiRNA和ΔCTS-LPAT1或AGPAT1的pBinGlyBar1骨架中(3)构建了多基因T-DNA载体。我们选择三个amiRNA序列靶向欧洲油菜LPAT2-样基因(5’-TCACTTGATGTGAAGATGCAC-3’(SEQ ID NO：30),5-’TTAACAGCTGACACGAAGCCT-3’(SEQ ID NO：31)和5’-TCACTTGATGTGAACACGCAC-3’(SEQ IDNO：32))，并将它们用于取代拟南芥MIR319a前体中的茎环(16)。采用根癌农杆菌将该构建体转化到欧洲油菜cv DH12075(WT)中(18)。获得了多个独立系的纯合T3种子批次，并如实施例1中所述开展脂质分析。当我们进行位置分析时，我们发现sn-2位C16：0的百分含量在WT种子中是～2％，在ProOLE：FATBProNAP：LPATamiRNA ProGLY：ΔCTS-LPAT1和ProOLE：FATBProNAP：LPATamiRNAProGLY：AGPAT种子中分别高达47％和62％(表9)。转基因系种子中的C16：0总含量也增加到24％至30％。数据表明，TAG中sn-2位的C16：0富集可以在经过修饰而提高C16：0总含量的欧洲油菜种子中产生。
FATB DNA序列(拟南芥)
FATB氨基酸序列(拟南芥)
表9.欧洲油菜WT种子、ProOLE：FATBProNAP：LPATamiRNA ProGLY：ΔCTS-LPAT1种
子和ProOLE：FATBProNAP：LPATamiRNA ProGLY：AGPAT1种子TAG中C16：0的总含量和sn-2位
C16：0的含量。数值是每种基因型三株植物的不同种子批次的测量平均值±SE。
实施例6-解脂耶氏酵母
我们在产油酵母解脂耶氏酵母中过表达ΔCTS-LPAT1(PLPAT，SEQ ID NO：38)、AGPAT1(SEQ ID NO：37)或莱茵衣藻(Chlamydomonas reinhardtii)LPAT(CRELPAT,SEQ IDNO：36)。当我们进行TAG位置分析时，我们发现在WT中，sn-2位C16：0的百分含量是～3％，在一些突变株中增加至高达～63％(表14)。数据表明，TAG中sn-2位的C16：0富集可以在解脂耶氏酵母细胞中产生。
本研究中使用的大肠杆菌菌株和酵母菌株分别列于表10和表11，Sambrook和Russell(2001)(20)以及Barth和Gaillardin(19)对它们的培养维持和生长条件分别进行了描述(1996)(19)。解脂耶氏酵母培养基、培养生长条件和在限氮下用于脂质提取的生物质收获与之前报道的相同(Bhutada et al.,2017)(23)。对于ura3Δ或leu2Δ营养缺陷型菌株的生长，培养基中补充0.1g L-1尿嘧啶或亮氨酸。
所有用于测序模板克隆和扩增的PCR反应均使用Herculase II Fusion DNAPolymerase(Agilent Technologies)和GoTaq(普洛麦格)进行，以确认转化盒的染色体整合。本研究中使用的限制酶来自Roche或New England Biolabs(NEB)。使用GeneJET试剂盒(赛默科技)从琼脂糖凝胶中回收来自PCR和限制性消化的DNA片段。对于连接，使用Fast-Link DNA连接试剂盒(Epicenter)或Gibson assembly(吉布森组装)(Gibson et al.,2009(21)；Kulasekara,2011(22))。为了转化为解脂耶氏酵母，使用醋酸锂标准方案(Le Dallet al.,1994(25))。所有引物都在表12中列出。
表10.本研究中使用的质粒。
表11.本研究中使用的菌株。
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表12.本研究中使用的引物
构建具有溶血磷脂酸酰基转移酶(LPAT)表达的突变体
为了获得具有LPAT表达的菌株，进行了属于人类、植物和藻类物种的密码子优化的LPAT CDS序列以及Tsynt25(SynT)合成终止子片段的基因合成(Curran et al.,2015)。先前描述了质粒pGMKGSY_12(Bhutada et al.,2017)，其含有两侧为糖原合酶1kb重组区的糖原贮积消除盒，GSY1位点采用HindIII消化线性化，并用于通过吉布森组装与TEF1启动子片段组装的LPAT基因的强组成型表达，TEF1启动子片段采用引物TEF-GSY-F/TEF-GSY-R从W29基因组DNA进行PCR扩增，得到质粒pGSYTEF。
采用HindIII消化含有LPAT CDS-SynT的基因合成pUC7载体以切除对应于AGPAT、CRELPAT和PLPAT的克隆插入物。用相同的酶消化pGSYTEF以使载体线性化，并在TEF1P启动子片段下，将其与上述凝胶纯化的插入物重新连接，产生pTEFAGPAT1、pTEFCRELPAT和pTEFPLPAT。通过测序，确定游离YlGSY1P-loxP-URA3-loxP-TEFPAGPAT1SynT-YlGSY1T、YlGSY1P-loxP-URA3-loxP-TEFPCRELPATSynT-YlGSY1T和YlGSY1P-loxP-URA3-loxP-TEFPPLPATSynT-YlGSY1T盒是否正确组装。这些盒经切下、NotI消化、纯化，用于菌株PO1d的转化(表11)。通过Lugol碘染色(1％KI,0.5％I2)鉴定在GSY1位点处与所述盒整合的转化体，并通过对照引物PCR和测序确认。
分析方法
如实施例1中所述提取和分析脂质。
表13.限氮培养基中解脂耶氏酵母菌株TAG的总脂肪酰基组成和sn-2脂肪酰基组
成。数值是每种基因型三种不同培养物的测量平均值±SD。
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表14.限氮培养基中解脂耶氏酵母菌株TAG中sn-2位的总C16：0富集。数值是每种
基因型三种不同培养物的测量平均值±SD。
用于解脂耶氏酵母中表达的优化基因序列合成参考下面的序列，下划线部分表示编码序列在每个末端添加限制性位点和3'UTR/终止子。
CreLPAT–
AGPAT1–
PLPAT–
实施例7—在各种碳源上培养解脂耶氏酵母
在实施例6中，评估了以20g L-1甘油作为碳源在限氮培养基上生长的解脂耶氏酵母的TAG脂肪酰基组成和sn-2位的C16：0百分含量。在本实施例中，我们还评估了使用糖、植物油和这些碳源的混合物作为碳源，考察它们对TAG中sn-2位的C16：0百分含量、TAG的总脂肪酰基组成、总脂质含量和生物质的形成是否有影响。本实施例中使用的糖、油和混合碳源如下：
20g L-1甘油(参见实施例6)
20g L-1葡萄糖
20g L-1棕榈油
10g L-1葡萄糖和10g L-1棕榈油
10g L-1甘油和10g L-1棕榈油
除了当细胞采用棕榈油培养时，培养基补充0.1％Tween-80/克所用油之外，解脂耶氏酵母菌株的培养基和生长条件与实施例6相同。如实施例1所述进行脂质提取和分析。
葡萄糖培养
表15和表19中提供的数据表明，当在限氮培养基中向解脂耶氏酵母WT、gsy1Δ和gsy1Δ-CreLPAT菌株提供葡萄糖作为碳源时，TAG脂肪酰基组成和sn-2位的％C16：0(表19)与在甘油上培养菌株时观察到的相似(实施例6)。
表15.以葡萄糖作为碳源时限氮培养基中解脂耶氏酵母菌株TAG的总脂肪酰基组
成和sn-2脂肪酰基组成。数值是每种基因型三种不同培养物的测量平均值±SD。nd是未检
测。
棕榈油培养
表16和表19中的数据表明，与甘油或葡萄糖作为碳源的培养物相比，当在限氮培养基中向解脂耶氏酵母WT和gsy1Δ-CreLPAT菌株提供棕榈油作为碳源时，TAG脂肪酰基组成发生改变。在gsy1Δ-CreLPAT中，TAG中C16：0的总百分含量从～20％增加到～26％，硬脂酸(C18：0)的总百分含量从～8％下降到～1％。对于gsy1Δ-CreLPAT，在棕榈油、葡萄糖或甘油上培养时，sn-2位的C16：0百分含量相似。然而，与在葡萄糖或甘油上培养相比，当gsy1Δ-CreLPAT在棕榈油上培养时，sn-1/3位的总长链饱和脂肪酰基(C16：0+C18：0)的百分含量降低，因为TAG中C18：0的总％降低，C18：0主要在sn-1/3位酯化。在WT和gsy1Δ-CreLPAT中，与葡萄糖或甘油上培养物相比，棕榈油上培养物的细胞生物量、细胞脂质含量和脂质滴度都更高(表20)。
表16.以棕榈油作为碳源时限氮培养基中解脂耶氏酵母菌株TAG的总脂肪酰基组
成和sn-2脂肪酰基组成。数值是每种基因型三种不同培养物的测量平均值±SD。nd是未检
测。
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在葡萄糖和棕榈油上培养
表17和表19中的数据表明，当在限氮培养基中向解脂耶氏酵母WT和gsy1Δ-CreLPAT菌株提供葡萄糖和棕榈油混合物作为碳源时，与仅采用葡萄糖或棕榈油作为碳源的培养物相比，TAG脂肪酰基组成发生改变。与葡萄糖或甘油培养物相比，gsy1Δ-CreLPAT中TAG内C16：0的总百分含量增加(至～24％)，C18：0的总百分含量亦降低。对于gsy1Δ-CreLPAT，与单独在葡萄糖、棕榈油或甘油上培养相比，sn-2位的C16：0百分含量更高。与单独在葡萄糖、棕榈油或甘油上培养相比，葡萄糖+棕榈油上培养时sn-1/3位的总长链饱和脂肪酸(C16：0+C18：0)的％更低。人乳脂(HMF)通常具有20％至25％的总％C16：0含量，在sn-2位具有～70％的C16：0和相对较低的～5％含量的总C18：0。因此，当在葡萄糖+棕榈油上培养时，gsy1Δ-CreLPAT的TAG脂肪酰基组成是更好的替代品，可提供足够的C16：0总百分含量，同时不仅降低了sn-1/3位的C16：0含量，还降低了总长链饱和脂肪酰基(C16：0+C18：0)的含量。在WT和gsy1Δ-CreLPAT中，与葡萄糖或甘油上培养物相比，葡萄糖+棕榈油上培养物的细胞生物量、细胞脂质含量和脂质滴度都更高(表20)。
表17.以棕榈油和葡萄糖作为碳源时限氮培养基中解脂耶氏酵母菌株TAG的总脂
肪酰基组成和sn-2脂肪酰基组成。数值是每种基因型三种不同培养物的测量平均值±SD。
nd是未检测。
在甘油和棕榈油上培养
表18、表19和表20中提供的数据表明，当在限氮培养基中向解脂耶氏酵母gsy1Δ-CreLPAT提供甘油和棕榈油的混合物作为碳源时，TAG脂肪酰基组成、sn-2位的C16：0％(表19)、细胞生物量、脂质含量和脂质滴度与在葡萄糖+棕榈油上培养的菌株时观察到的更相似，而与单独在葡萄糖或甘油上培养的相似程度更低些。在实施例6和7中使用的碳源上培养的解脂耶氏酵母WT或gsy1Δ-CreLPAT中未检测到二棕榈酰PC。使用实施例1中描述的方法分析样品。这表明表达C16：0LPAT的解脂耶氏酵母细胞将C16：0从PC的sn-2位排除；我们在表达C16：0LPAT的拟南芥种子中也观察到了这一点。因此，在表达C16：0LPAT的解脂耶氏酵母中C16：0结合到TAG的sn-2位也可能受到负责DG/PC转化的酶活性的限制，例如C16：0结合到TAG的sn-2位可以通过抑制或阻止胆碱磷酸转移酶(CPT1；EC 2.7.8.2)和/或乙醇胺磷酸转移酶(EPT1；EC 2.7.8.1)的活性来增强。
表18.以棕榈油和甘油混合物作为碳源时限氮培养基中解脂耶氏酵母菌株TAG的
总脂肪酰基组成和sn-2脂肪酰基组成。数值是每种基因型单个培养物的测定值。
表19.在不同碳源的限氮培养基中解脂耶氏酵母菌株TAG中sn-2位的总C16：0富
集。数值是每种基因型三种不同培养物的测定平均值±SD，但甘油+棕榈油上gsy1Δ-
CreLPAT的数值是单种培养物的测定值。
表20.在不同碳源的限氮培养基中解脂耶氏酵母菌株的生物量、脂质含量和脂质 滴度。数值是每种基因型三种不同培养物的测定平均值±SD，但甘油+棕榈油上gsy1Δ- CreLPAT的数值是单种培养物的测定值。
实施例8
尽管上面讨论的实施例表明，植物脂质代谢可以经工程改造使16：0优先酯化到TAG中的sn-2位，但希望拟南芥种子的总脂肪酸组成更接近人乳的总脂肪酸组成。拟南芥种子中16：0的含量低约3倍，并且它们含有高比例的多不饱和脂肪酸和超长链脂肪酸，而这些脂肪酸在人乳中基本不存在。人乳中最丰富的脂肪酸是18：1，并且由于次丰富的脂肪酸16：0不寻常的区域特异性分布，TAG的主要分子种类通常是1,3-二油酸-2-棕榈酸甘油三酯(OPO)，占总含量的～14％。我们决定通过将TAG中sn-2位的16：0富集与富含适当比例的16：0和18：1的总脂肪酸组成相结合，研究拟南芥种子是否可以经工程改造产生OPO。
结果
fab1-1 fae1fad2 fab1-1 fae1fad2种子中16：0和18：1含量均较高(HPHO)
要获得16：0含量相当于人乳的拟南芥种子，必须将其含量提高至～3倍达到20-25％。实现这一目标的一种方法是通过破坏β-酮脂酰-ACP合酶II基因脂肪酸生物合成1(FAB1)来减少脂肪酸合酶催化的16：0延伸(图8)。FAB1是拟南芥的必需基因，但已经表征了其种子中含～17％16：0的亚效fab1-1等位基因。然后在缺乏极长链脂肪酸合成的脂肪酸延长酶1(fae1)突变背景中，16：0可以增加到～24％(图8)。但是，fab1-1 fae1种子仍然含有比例较高的通过脂肪酸去饱和酶2(FAD2)产生的多不饱和脂肪酸(图8)。因此，为了创建16：0和18：1均高(HPHO)的背景，我们通过杂交构建了fab1-1 fae1 fad2突变体。对纯合fab1-1fae1fad2种子的分析表明，TAG的脂肪酸组成中16：0和18：1均高，分别占总脂肪酸的～20％和～70％(图9)。野生型拟南芥Col-0种子TAG中通常含量丰富的其他脂肪酸种类，如亚油酸(18：2)、亚麻酸(18：3)和二十碳烯酸(20：1)，在此纯合种子中各自含量均<3％(图9)。与双突变体的比较表明，fad2背景中fab1-1 fae1中的16：0含量显著降低(P>0.05)(图9)。但是，fab1-1 fae1fad2种子的HPHO组成表明这种遗传背景适合试验种子中是否可以产生OPO。
HPHO种子中的ΔCTS-LPAT1表达促进16：0结合到TAG的sn-2位
如上所述，我们已经表明，在大豆球蛋白-1启动子(ProGLY)下，在WT拟南芥种子中表达ER重新靶向的叶绿体LPAT(ΔCTS-LPAT1)会导致TAG中sn-2位酯化的16：0显著增加。为了确定ΔCTS-LPAT1表达在HPHO背景中的影响，我们通过杂交构建了ProGLY：ΔCTS-LPAT1fab1-1fae1 fad2系。当我们从种子中纯化TAG并测定其区域化学时，我们发现sn-2位(与sn-1+3位相比)16：0的百分含量从fab1-1 fad2 fae1背景中的～3％增加到ProGLY：ΔCTS-LPAT1 fab1-1fad2 fae1的～24％(图10A)。除16：0丰度发生虽小但显著(P>0.05)增加，从～19％增加到～22％之外，fab1-1 fad2 fae1种子的总脂肪酸组成并没有因ΔCTS-LPAT1表达而发生很大变化(图10B)。我们之前观察到，当我们在WT种子中表达ΔCTS-LPAT1时，总16：0含量略有增加。我们的数据表明，HPHO种子中的ΔCTS-LPAT1表达允许16：0结合到TAG的sn-2位。然而，富集水平显著(P>0.05)低于含ProGLY：ΔCTS-LPAT1的WT种子(图10A)。
AGPAT1表达促进C16：0更强地结合到TAG的sn-2位
为了研究与ΔCTS-LPAT1相比，其他ER定位的LPAT是否可能使拟南芥将更多的16：0结合到TAG的sn-2位，我们决定测试人类AGPAT1。人乳脂肪球由乳腺上皮中的乳腺细胞分泌。目前尚不清楚哪种LPAT负责人乳脂的生物合成。然而，AGPAT1在乳腺上皮细胞中表达，体外试验表明AGPAT1可以使用16：0-CoA作为底物。为了测试AGPAT1是否可以将16：0结合到TAG的sn-2位，我们首先在GAL1启动子下在酿酒酵母中表达该蛋白质(28)。当我们纯化TAG，并测定其区域化学时，我们发现sn-2位16：0的百分含量从含有空白载体对照的细胞中的～4％增加到含有ProGAL1：AGPAT1的细胞中的～45％(表21)。
表21.表达AGPAT1的酿酒酵母细胞中TAG的16：0
数值是三种不同培养物的脂质提取物的测量平均值±SE。*表示与空白载体对照(EVC)(双尾学生t-检验)显著不同的数值(P<0.05)。
使用本氏烟叶中的瞬时表达，我们还证明：当AGPAT1在花椰菜花叶病毒35S启动子下以红色荧光蛋白(RFP)-AGPAT1融合蛋白表达时，它也可以定位到植物细胞中的ER(图11A)。接下来，我们用ProGLY：AGPAT1构建体转化WT拟南芥植物，以促进转基因的强种子特异性表达。我们从>40个初级转化体中选择两个独立的单拷贝T2系(L35和L40)进行分析，并获得纯合T3种子。当我们从纯合种子批次中纯化TAG并测定其区域化学时，我们发现，L40的sn-2位16：0的百分含量是～66％，L35是～74％(图11B)。为了确定AGPAT1表达在HPHO背景中的影响，我们通过杂交构建了ProGLY：AGPAT1 fab1-1fae1 fad2系。当我们从这些种子纯化TAG并测定其区域化学时，我们发现，sn-2位16：0的百分含量是～54％(图11B)。ProGLY：AGPAT1 fab1-1 fae1 fad2种子的总16：0含量与fab1-1 fae1 fad2并未显著(P>0.05)不同(图11C)。因此，与ΔCTS-LPAT1相比，AGPAT1表达允许将更高含量的16：0结合到WT种子和HPHO种子中TAG的sn-2位。
破坏LPAT2和PDCT增强16：0的ΔCTS-LPAT1和AGPAT1-依赖性结合到HPHO种子内TAG的sn-2位
在野生型(WT)拟南芥种子中，我们之前发现通过破坏酶LPAT2和PDCT可以增加16：0的ΔCTS-LPAT1依赖性结合到TAG的sn-2位。LPAT2是拟南芥种子中表达的主要的ER定位LPAT同种型，因此其破坏可能会减少与ΔCTS-LPAT1的竞争(图8)。PDCT催化拟南芥种子中的快速DG/PC相互转化。尽管LPAT最初在sn-2位酰化甘油脂，但一旦这些酰基在PC中，它们就可以通过脱酰-再酰化(酰基编辑)循环删除和取代。PDCT的破坏迫使新制造的DG更直接地进入TAG(图8)。为了确定LPAT2和PDCT破坏是否影响表达ΔCTS-LPAT1或AGPAT1的HPHO种子中酯化到TAG中sn-2位的16：0的百分含量，我们通过杂交构建了ProGLY：ΔCTS-LPAT1fab1-1 fae1fad2 lpat2-2 pdct和ProGLY：AGPAT1 fab1-1 fae1 fad2lpat2-2 pdct系。当我们从这些种子纯化TAG并测定其区域化学时，我们发现，sn-2位16：0的百分含量分别是～62％和～83％(图12A)。ProGLY：ΔCTS-LPAT1 fab1-1 fae1 fad2 lpat2-2 pdct和ProGLY：AGPAT1 fab1-1 fae1 fad2 lpat2-2 pdct种子中的总16：0含量分别是～21和～18％(图12B)。
HPHO种子的含油量和种子活力降低，但这种情况并不会因16：0重新分布到sn-2位而加剧。
修饰脂肪酸组成会减少油籽中TAG的积累，在某些情况下还会损害种子发芽和成苗。如上所述，我们发现，16：0总含量低，但～70％酯化到sn-2位的ProGLY：ΔCTS-LPAT1lpat2-3pdct种子显示出TAG占种子重量的百分含量降低。然而，它们的发芽和初始幼苗生长并没有明显受到损害。为了考察HPHO种子中16：0富集在TAG的sn-2位的生理影响，我们对在标准实验室条件下一起生长的WT、fab1-1 fae1fad2、ProGLY：ΔCTS-LPAT1 fab1-1fae1fad2 lpat2-2 pdct和ProGLY：AGPAT1 fab1-1 fae1fad2 lpat2-2 pdct植物的种子批次进行了比较。我们发现，与WT相比，fab1-1 fae1 fad2的种子重量和含油量显著降低(P>0.05)(图13)。但是，fab1-1 fae1 fad2和ProGLY：ΔCTS-LPAT1 fab1-1 fae1fad2 lpat2-2pdct或ProGLY：AGPAT1 fab1-1 fae1fad2 lpat2-2 pdct种子之间并未观察到任何显著差异(图13)。这些数据表明，fab1-1 fae1 fad2种子中的TAG生物合成通量受到损害，但导致16：0结合到TAG的sn-2位的进一步遗传修饰在此背景下并不存在损害。这些发现与我们在WT种子中观察到的结果形成对比。在标准实验室条件下(20℃，16h光周期)，fab1-1fae1fad2种子萌发(以胚根出现计分)和幼苗成苗(以子叶展开计分)的速度显著(P>0.05)低于WT(图14)。但是，fab1-1 fae1 fad2和ProGLY：ΔCTS-LPAT1 fab1-1 fae1fad2 lpat2-2 pdct或ProGLY：AGPAT1 fab1-1 fae1fad2 lpat2-2 pdct种子之间并未观察到任何显著差异(P>0.05)(图14)。
讨论
在本实施例中，我们表明了拟南芥种子可以经工程改造产生OPO，因为通过结合fab1-1、fae1和fad2等位基因，TAG含有～20％的16：0和～70％的18：1，其中>80％的16：0被酯化到甘油骨架的sn-2位。OPO是人乳中的主要TAG种类，但它几乎不存在于植物油中，植物油中酯化到sn-2位的16：0的含量非常低。人乳中的高OPO含量被认为具有营养价值，因此开发富含OPO的植物油可以为婴儿配方奶粉成分提供有用的新来源。
WT拟南芥种子中重新靶向的叶绿体LPAT(ΔCTS-LPAT1)的表达允许TAG中～30％至40％的16：0占据sn-2位。然而，当我们在HPHO背景中表达ΔCTS-LPAT1时，我们发现结合到sn-2位的16：0降低至～20％。然后，LPAT2和PDCT的破坏导致sn-2位16：0的百分含量增加到～62％。sn-2位16：0的这种富集水平亦低于我们使用相同方法在WT中所能达到的水平。然而，HPHO的总脂肪酸组成更适合婴儿配方奶粉成分，并且sn-2位～62％16：0仍优于通过体外酶催化生产的市售HMFS。
通常，在人乳脂中，>70％的16：0被酯化到TAG的sn-2位，因此这种富集水平仍然是HMFS的目标。人LPAT AGPAT1可以使用16：0-CoA作为底物并在乳腺细胞中表达。当我们在WT拟南芥种子中表达AGPAT1时，我们发现，TAG中60％到70％的16：0占据sn-2位。
当我们在HPHO背景中表达AGPAT1时，结合到sn-2位的16：0减少到～54％，但LPAT2和PDCT的破坏随后导致sn-2位16：0的百分含量增加到～83％。sn-2位16：0的这种富集水平大于或等于人乳脂中报告的水平。
代谢途径工程通常会对油籽中TAG的积累产生不利影响，并可能损害种子活力。在上面讨论的实施例1中，我们发现在WT拟南芥种子中将16：0重新定位到TAG的sn-2位减少了油的积累。HPHO种子的种子含油量也低于WT。然而，在16：0的位置分布中设计类似的变化并没有导致进一步的减少。鉴于WT种子的16：0含量比HPHO种子低～3倍，可能是16：0的可利用性限制了经工程改造仅具有16：0-CoA LPAT活性的种子中TAG生物合成的速率。相反，HPHO种子中16：0含量高～3倍可能会限制TAG生物合成，因为天然LPAT(和其他酰基转移酶活性)的16：0-CoA活性太低。HPHO种子在种子发芽和早期幼苗生长方面也受到明显损害。然而，将16：0重新定位到HPHO种子中TAG的sn-2位并未加剧这种影响。HPHO种子活力更差可能是由于长链脂肪酸不饱和度降低，从而提高了油的熔化温度。这种性质受脂肪酸位置分布的影响不是很大。
材料和方法
植物材料和生长条件
以前已对拟南芥哥伦比亚-0(Colombia-0)突变体fab1-1、fae1、fad2、pdct和lpat2-3进行了描述(26)(27)(2)。对于在培养基上进行的实验，对来自单株植物的～50颗种子进行表面消毒，将其铺在含有一半强度Murashige和Skoog盐(Sigma-Aldrich)、pH 5.7的琼脂板上，并在4℃下在黑暗中吸胀4天。然后将琼脂板放置在设置为16小时光照(光合光子通量密度＝150μmol m-2s-1)/8小时黑暗周期、恒温20℃的生长室中。如前文所述，在解剖立体显微镜下对发芽(胚根出现)和子叶展开进行视觉评分。还将单株幼苗移植到含有Levington F2堆肥的7cm2盆中，并在设置为16小时光照(22℃)/8小时黑暗(16℃)周期，其中光合光子通量密度是250μmol m-2s-1的室中生长。在开花开始时将植物单独套袋，并在成熟时收获种子。
基因分型
使用DNeasy Plant Mini Kit(Qiagen)从叶子分离基因组DNA。如前文所述，使用普洛麦格PCR Master Mix(普洛麦格)和基因特异性和T-DNA左边界引物对的组合通过PCR鉴定纯合lpat2-3T-DNA插入突变体。纯合fab1-1、fad2、fae1和pdct突变体通过对采用跨越点突变位点的引物对扩增的PCR产物进行测序来鉴定(26)(27)(2)。如前所述，ProGLY：ΔCTS-LPAT和ProGLY：AGPAT1T-DNA的存在通过使用跨越ProGLY和ΔCTS-LPAT1或AGPAT1的引物对的PCR确定。
脂质分析
从材料中提取总脂质，并如前所述纯化TAG(6)。除了2-单酰基甘油使用己烷：乙醚：乙酸(35：70：1.5,v/v/v)通过薄层色谱法(硅胶60,20 x 20cm；Sigma-Aldrich/Merck)(8)分离之外，TAG区域化学分析按照前面描述的方法通过脂肪酶消化进行。如前所述，使用配备DB-23柱(30mx 0.25mm i.d.x 0.25μm)的7890A GC系统(安捷伦科技公司)，通过连接火焰离子化检测的气相色谱(GC)，对整个种子和纯化脂质中存在的脂肪酰基进行反式甲基化和定量。如前所述，使用配备机器人样品处理系统(Rohasys)的Minispec MQ20设备(Bruker)，通过低分辨率时域NMR光谱测量整个种子的种子含油量和水分含量，并将含油量百分数标准化为9％水分。
克隆和转化
人类AGPAT1(GenBank：NP_001358367,SEQ ID NO：40)经密码子优化用于在拟南芥中表达，其由Genscript合成并以pUC57提供。然后使用引物对5’-CACCATGGATTTATGGCCTGGTGC-3’(SEQ ID NO：74)&5'-TCATCCTCCTCCACCTGG-3'(SEQ ID NO：75)，采用KOD DNA聚合酶(Merck)通过PCR扩增AGPAT1。使用QIAquick凝胶回收试剂盒(Qiagen)纯化得到的PCR产物。PCR产物在pENTR/D-TOPO载体(赛默飞世尔科技)中克隆，测序(SEQ ID NO：26)，并使用Gateway LR Clonase II Enzyme mix(赛默飞世尔科技)重组到pYES-DEST52(Invitrogen)和pK7WGR2中。AGPAT被克隆到位于大豆球蛋白-1(GLY)启动子和种子特异性表达终止子之间的pBinGlyRed3载体中(3)。AGPAT1使用KOD DNA聚合酶和引物对5'-CGGAATTCATGGATTTATGGCCTGGTGC-3'(SEQ ID NO：76)&5'-GCTCTAGATCATCCTCCTCCACCTGG-3'(SEQ ID NO：77)从pENTR-D-TOPO载体开展PCR扩增。PCR产物经凝胶纯化并用EcoRI和XbaI消化。pBinGlyRed3载体也用EcoRI和XbaI消化，碱性磷酸酶处理(普洛麦格)，凝胶纯化，并使用T4 DNA连接酶(NEB)将AGPAT1连接到该载体中。使用S.c.EasyComp试剂盒(Invitrogen)将pYES-DEST52转化为酿酒酵母INVSc1细胞，并如Kimet al.,(2005)(28)所述诱导蛋白质表达。使用热激将pK7WGR2和pBinGlyRed3载体转化到根癌农杆菌菌株GV3101内。拟南芥转化使用浸花法(4)进行。使用DsRed滤光片在LeicaM205 FA显微镜下鉴定表达可选择标记的T1种子。
本氏烟中的瞬时表达和成像
正如Wood et al.,(2009)(5)所述，采用含Pro35S：RFP-AGPAT1,Pro35S：m-GFP5-ER或Pro35S：p19的载体转化的根癌农杆菌培养物在本氏烟叶中进行瞬时表达。将培养物手工渗透到叶子中，并将接种的植物放置48小时。然后在Apochromat 63x/1.20W Korr M27物镜下，将本氏烟叶安装在水中置于Zeiss LSM 780激光扫描共聚焦显微镜上。GFP在488nm的波长处被激发，RFP在561nm处被激发。检测采用具有473-551nm发射带的滤光片。
统计分析
所有实验均使用三个生物重复进行，数据以平均值±平均值标准误差(SE)表示。对于统计分析，我们使用单向方差分析(ANOVA)和事后Tukey HSD(真实显著性差异)检验，或使用双尾学生t-检验。
登陆号
本文所述的序列数据见GenBank/EMBL数据库中例如登录号：NP_001358367(AGPAT1)、AF111161(LPAT1)、At1g74960(FAB1)、At3g12120(FAD2)、At4g34520(FAE1)、At3g57650(LPAT2)、At3g15820(PDCT)。
附录–附加序列信息
>HsAGPAT1 cDNA序列
>HsAGPAT1氨基酸序列
>CreLPAT cDNA序列(莱茵衣藻基因Cre17.g738350)
/>
>CreLPAT氨基酸序列
>LC107611.1微拟球藻属NIES-2145 LPAT2 mRNA，用于溶血磷脂酰转移酶2，完整的蛋白质编码区
>微拟球藻属NIES-2145 LPAT2氨基酸序列
>LC107612.1微拟球藻属NIES-2145 LPAT3 mRNA，用于溶血磷脂酰转移酶3，完整的蛋白质编码区
>微拟球藻属NIES-2145 LPAT3氨基酸序列
>LC107613.1微拟球藻属NIES-2145 LPAT4 mRNA，用于溶血磷脂酰转移酶4，完整的蛋白质编码区
>微拟球藻属NIES-2145 LPAT4氨基酸序列
>BAA18602.1 sll1848[集胞藻属PCC 6803]mRNA序列
>BAA18602.1 sll1848[集胞藻属PCC 6803]氨基酸序列
编码拟南芥ER LPAT2的核苷酸序列：
>ENA|AEE79683|AEE79683.1拟南芥(thale cress)溶血磷脂酰转移酶2
拟南芥ER LPAT2的氨基酸序列：
>sp|Q8LG50|LPAT2_ARATH 1-酰基-sn-甘油-3-磷酸酰基转移酶2Os＝拟南芥OX＝3702 GN＝LPAT2 PE＝1 SV＝2
编码向日葵ER LPAT2的核苷酸序列：
>ENA|OTG36509|OTG36509.1向日葵(普通向日葵)推定的1-酰基-sn-甘油-3-磷酸酰基转移酶2
向日葵LPAT2的氨基酸序列：
>tr|E6Y2I1|E6Y2I1_HELAN推定的1-酰基-sn-甘油-3-磷酸酰基转移酶2 OS＝向日葵OX＝4232 GN＝LPAAT2 PE＝2 SV＝1
编码亚麻荠ER LPAT2的核苷酸序列：
>XM_010429345.2预测：亚麻荠1-酰基-sn-甘油-3-磷酸酰基转移酶2(LOC104712441),mRNA
亚麻荠ER LPAT2的氨基酸序列：
>XP_010427647.1预测：1-酰基-sn-甘油-3-磷酸酰基转移酶2[亚麻荠]
编码欧洲油菜ER LPAT2的核苷酸序列：
>ENA|CAB09138|CAB09138.1欧洲油菜(油菜)酰基辅酶A：1-酰基甘油-3-磷酸酰基转移酶
欧洲油菜ER LPAT2的氨基酸序列：
>sp|Q9XFW4|LPAT2_BRANA 1-酰基-sn-甘油-3-磷酸酰基转移酶2 Os＝欧洲油菜OX＝3708 GN＝LPAT2 PE＝2 SV＝1
编码大豆ER LPAT2的核苷酸序列：
>XM_003535809.4大豆1-酰基-sn-甘油-3-磷酸酰基转移酶2(LOC100814107),mRNA
大豆LPAT2的氨基酸序列：
>XP_003535857.1 1-酰基-sn-甘油-3-磷酸酰基转移酶2[大豆]
编码解脂耶氏酵母ER LPAT的核苷酸序列：
>ENA|CAG79722|CAG79722.1解脂耶氏酵母CLIB122 YALI0E18964p：位置：1..849
解脂耶氏酵母ER LPAT的氨基酸序列：
>tr|Q6C5D5|Q6C5D5_YARLI 1-酰基-sn-甘油-3-磷酸酰基转移酶OS＝解脂耶氏酵母(菌株CLIB 122/E 150)OX＝284591 GN＝YALI0_E18964g PE＝3 SV＝1
编码拟南芥PDCT的核苷酸序列：
>ENA|AEE75730|AEE75730.1拟南芥(thale cress)磷脂酸磷酸酶相关/PAP2样蛋白
拟南芥PDCT的氨基酸序列：
>sp|Q9LVZ7|PDCT1_ARATH磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1 Os＝拟南芥OX＝3702 GN＝ROD1 PE＝1 SV＝1
编码向日葵PDCT的核苷酸序列：
>ENA|ARQ87993|ARQ87993.1向日葵(普通向日葵)磷脂酰胆碱：位置：1..840
编码向日葵PDCT的核苷酸序列：
>tr|A0A3S5FWJ3|A0A3S5FWJ3_HELAN磷脂酰胆碱OS＝向日葵 OX＝4232 GN＝PDCTPE＝2 SV＝1
编码亚麻荠PDCT的核苷酸序列：
>XM_010467272.2预测：亚麻荠磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1(LOC104745897),mRNA
亚麻荠PDCT的氨基酸序列：
>XP_010465574.1预测：磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1[亚麻荠]
编码欧洲油菜PDCT的核苷酸序列：
>XM_013791764.2预测：欧洲油菜磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1(LOC106352083),mRNA
欧洲油菜PDCT的氨基酸序列：
>XP_013647218.1磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1[欧洲油菜]
编码大豆PDCT的核苷酸序列：
>XM_003531670.4预测：大豆磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1(LOC100790665),mRNA
大豆PDCT的氨基酸序列：
>XP_003531718.1磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶1[大豆]
拟南芥At1g74960(FAB1)核苷酸序列
拟南芥At1g74960(FAB1)氨基酸序列
拟南芥At3g12120(FAD2)核苷酸序列
拟南芥At3g12120(FAD2)氨基酸序列
拟南芥At4g34520(FAE1)核苷酸序列
拟南芥At4g34520(FAE1)氨基酸序列
解脂耶氏酵母YALI0B10153g(FAD2)核苷酸序列
解脂耶氏酵母YALI0B10153p(FAD2)氨基酸序列
解脂耶氏酵母YALI0B20196p(FAE1/ELO2)核苷酸序列
解脂耶氏酵母YALI0B20196p(FAE1/ELO2)氨基酸序列
解脂耶氏酵母YALI0C10989g(CPT1)核苷酸序列
解脂耶氏酵母YALI0C10989p(CPT1)氨基酸序列
解脂耶氏酵母YALI0E26565g(EPT1)核苷酸序列
解脂耶氏酵母YALI0E26565p(EPT1)氨基酸序列
参考文献
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如本文所述，在人乳脂中，饱和脂肪酸酯化到甘油骨架的中间位置，使甘油三酯分子具有不寻常的立体化学，有助于婴儿肠道中的营养吸收。然而，大多数婴儿配方奶粉中使用的脂肪来自饱和脂肪酸酯化到外部位置的植物。在这里，我们设计了一种油料植物的新陈代谢，使其积聚中间位置具有超过70％的饱和脂肪酸棕榈酸酯的甘油三酯，从而模仿人乳脂的立体异构结构。将这项技术应用于油料作物(或产油微生物)可以为婴儿营养提供一种新的人乳脂替代品来源。
应该理解的是，对于本领域技术人员来说，对本文所述的当前优选实施例的各种改变和修改将是显而易见的。这些改变和修改可以在不背离本发明的精神和范围并且不削弱其附带优点的情况下进行。因此，这些变化和修改由所附权利要求覆盖。
本文引用的所有参考文献的内容均通过引用的方式全文并入本发明中。
序列表
<110> 罗特哈姆斯泰德研究有限公司
<120> 产生甘油三酯的非人类生物体
<130> P16317S-PCT
<160> 91
<170> PatentIn 版本 3.5
<210> 1
<211> 777
<212> DNA
<213> 拟南芥
<400> 1
atgtcggatc tttcaggagc tgcaacccct gaatctactt atccagaacc agagattaag 60
ttgagctcaa gactcagagg gatatgcttt tgtctcgttg ctggcgtttc cgccattgtt 120
ctcatcgtcc tgatgatcac tggccatcct ttcgtccttc tatttgatcg ttacaggagg 180
aagttccatc acttcattgc taagctctgg gcttccataa gcatctaccc gttttacaaa 240
accgacatcc aaggtttgga gaatctgcca tcatcagaca ctccttgtgt gtacgtttcg 300
aaccaccaga gttttctgga tatatacaca cttctcagcc ttggccaaag ctataagttc 360
atcagcaaga cagggatatt cgttattcct gtcatcggtt gggctatgtc catgatgggg 420
gttgttccct tgaagaggat ggacccaaga agccaagtgg attgcttaaa acgctgcatg 480
gaactagtga agaagggagc ttccgtcttt ttcttcccag agggaacgag gagtaaggat 540
ggtcggttag gtcctttcaa gaaaggggct tttacgatag cagctaagac aggagttcca 600
gtggtgccaa taacgctgat gggaacaggg aagatcatgc cgacgggtag tgaaggtata 660
ctgaatcatg gggatgtgag agtgatcatc cacaagccga tatatggaag caaagctgat 720
cttctttgcg atgaggctag aaacaagata gctgaatcta tgaatctctt gagttga 777
<210> 2
<211> 258
<212> PRT
<213> 拟南芥
<400> 2
Met Ser Asp Leu Ser Gly Ala Ala Thr Pro Glu Ser Thr Tyr Pro Glu
1 5 10 15
Pro Glu Ile Lys Leu Ser Ser Arg Leu Arg Gly Ile Cys Phe Cys Leu
20 25 30
Val Ala Gly Val Ser Ala Ile Val Leu Ile Val Leu Met Ile Thr Gly
35 40 45
His Pro Phe Val Leu Leu Phe Asp Arg Tyr Arg Arg Lys Phe His His
50 55 60
Phe Ile Ala Lys Leu Trp Ala Ser Ile Ser Ile Tyr Pro Phe Tyr Lys
65 70 75 80
Thr Asp Ile Gln Gly Leu Glu Asn Leu Pro Ser Ser Asp Thr Pro Cys
85 90 95
Val Tyr Val Ser Asn His Gln Ser Phe Leu Asp Ile Tyr Thr Leu Leu
100 105 110
Ser Leu Gly Gln Ser Tyr Lys Phe Ile Ser Lys Thr Gly Ile Phe Val
115 120 125
Ile Pro Val Ile Gly Trp Ala Met Ser Met Met Gly Val Val Pro Leu
130 135 140
Lys Arg Met Asp Pro Arg Ser Gln Val Asp Cys Leu Lys Arg Cys Met
145 150 155 160
Glu Leu Val Lys Lys Gly Ala Ser Val Phe Phe Phe Pro Glu Gly Thr
165 170 175
Arg Ser Lys Asp Gly Arg Leu Gly Pro Phe Lys Lys Gly Ala Phe Thr
180 185 190
Ile Ala Ala Lys Thr Gly Val Pro Val Val Pro Ile Thr Leu Met Gly
195 200 205
Thr Gly Lys Ile Met Pro Thr Gly Ser Glu Gly Ile Leu Asn His Gly
210 215 220
Asp Val Arg Val Ile Ile His Lys Pro Ile Tyr Gly Ser Lys Ala Asp
225 230 235 240
Leu Leu Cys Asp Glu Ala Arg Asn Lys Ile Ala Glu Ser Met Asn Leu
245 250 255
Leu Ser
<210> 3
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物 P1
<400> 3
caccatgtcg gatctttcag gagctg 26
<210> 4
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物 P2
<400> 4
tcaactcaag agattcatag attca 25
<210> 5
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物 P3
<400> 5
cggaattcat gtcggatctt tcaggagc 28
<210> 6
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物 P4
<400> 6
gctctagatc aactcaagag attcatagat tc 32
<210> 7
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物 P5
<400> 7
gttgattgaa tcgaggaagg a 21
<210> 8
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物 P6
<400> 8
ctttttacca catgcaaagg g 21
<210> 9
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P7
<400> 9
ccattttggt ggtcgtctat 20
<210> 10
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物 P8
<400> 10
cagaaaaatt aaccgggtgg t 21
<210> 11
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P9
<400> 11
cagctgaaac cgacgtctct 20
<210> 12
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P10
<400> 12
ggtcacgcgc tctttgaatc 20
<210> 13
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> 引物 P11
<400> 13
gtagcaccca aggcttcca 19
<210> 14
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> 引物 P12
<400> 14
gccagtgatc atcaggacg 19
<210> 15
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物 P13
<400> 15
gcgtggaccg cttgctgcaa ct 22
<210> 16
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P14
<400> 16
ctggtgggct ggagttaaga 20
<210> 17
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P15
<400> 17
agaatccatc ccacaagcca 20
<210> 18
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P16
<400> 18
tgtcatcggt tgggctatgt 20
<210> 19
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物 P17
<400> 19
catcagcgtt attggcacca 20
<210> 20
<211> 17
<212> DNA
<213> 人工序列
<220>
<223> 引物 P18
<400> 20
gacgcttcat ctcgtcc 17
<210> 21
<211> 16
<212> DNA
<213> 人工序列
<220>
<223> 引物 P19
<400> 21
ccacaggttg cgttag 16
<210> 22
<211> 16
<212> DNA
<213> 人工序列
<220>
<223> 引物 P20
<400> 22
tccagctaag ggtgcc 16
<210> 23
<211> 16
<212> DNA
<213> 人工序列
<220>
<223> 引物 P21
<400> 23
ggtgggtact cggaga 16
<210> 24
<211> 17
<212> DNA
<213> 人工序列
<220>
<223> 引物 P22
<400> 24
gaattacccg acggaca 17
<210> 25
<211> 16
<212> DNA
<213> 人工序列
<220>
<223> 引物 P23
<400> 25
acggtctgca atacct 16
<210> 26
<211> 852
<212> DNA
<213> 智人
<400> 26
atggatttat ggcctggtgc ttggatgtta ttattattat tattcttgtt attgctcttc 60
ctccttccta ctttgtggtt ctgttcacct tctgcaaagt atttctttaa gatggctttc 120
tacaacggat ggattctttt cttggctgtt ttggcaatcc cagtttgtgc tgtgagaggt 180
aggaacgttg aaaacatgaa gattctcaga ctcatgcttt tgcatatcaa gtacctttac 240
ggaataagag tggaagttag gggtgctcat cactttcctc catctcaacc ttatgttgtg 300
gtttcaaacc accagtcttc attggatctc ttaggaatga tggaagtgct ccctggtaga 360
tgtgttccaa tagcaaagag ggagcttttg tgggctggat cagcaggtct tgcttgctgg 420
ttggctggag ttatttttat cgatagaaaa aggacaggtg atgcaatctc agtgatgagt 480
gaagttgctc aaactctctt aacacaggat gtgagagttt gggtgttccc tgagggaacc 540
aggaatcata acggtagtat gttaccattt aagagaggag ctttccacct cgcagttcaa 600
gctcaggttc ctatagtgcc aatagttatg agttcttacc aagatttcta ctgtaagaag 660
gaaagaaggt tcactagtgg acaatgccaa gttagagtgc tccctccagt tccaaccgag 720
ggtttaactc ctgatgatgt gccagctctt gcagatagag ttaggcattc tatgttgaca 780
gtgttcagag agattagtac cgatggaagg ggaggtggag attatttgaa gaagccaggt 840
ggaggaggat ga 852
<210> 27
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 1
<400> 27
taaagcgagt tccctcgaca g 21
<210> 28
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 2
<400> 28
ttgtgcccag tgtacggact t 21
<210> 29
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 3
<400> 29
tcaaaggcac gatgatacct g 21
<210> 30
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 4
<400> 30
tcacttgatg tgaagatgca c 21
<210> 31
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 5
<400> 31
ttaacagctg acacgaagcc t 21
<210> 32
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> amiRNA 序列 6
<400> 32
tcacttgatg tgaacacgca c 21
<210> 33
<211> 1239
<212> DNA
<213> 拟南芥
<400> 33
atggtggcca cctctgctac gtcgtcattc tttcctgtac catcttcttc acttgatcct 60
aatggaaaag gcaataagat tgggtctacg aatcttgctg gactcaattc tgcacctaac 120
tctggtagga tgaaggttaa accaaacgct caggctccac ctaagattaa tgggaaaaag 180
gttggtttgc ctggttctgt agatattgta aggactgata ccgagacctc atcacaccct 240
gcgccgagaa ctttcatcaa ccagttacct gactggagca tgcttcttgc tgctataact 300
acgattttct tagcggctga gaaacagtgg atgatgcttg attggaaacc taggcgttct 360
gacatgctgg tggatccttt tggtataggg agaattgttc aggatggcct tgtgttccgt 420
cagaattttt ctattaggtc atatgaaata ggtgctgatc gctctgcatc tatagaaacc 480
gtcatgaatc atctgcagga aacggcgctt aatcatgtta agactgctgg attgcttgga 540
gatgggtttg gctctacacc tgagatgttt aagaagaact tgatatgggt tgtcactcgt 600
atgcaggttg tggttgataa atatcctact tggggagatg ttgttgaagt agacacctgg 660
gtcagtcagt ctggaaagaa tggtatgcgt cgtgattggc tagttcggga ctgtaatact 720
ggagaaacct taacacgagc atcaagtgtg tgggtgatga tgaataaact gacaaggaga 780
ttgtcaaaga ttcctgaaga ggttcgaggg gaaatagagc cttattttgt gaattctgat 840
cctgtccttg ccgaggacag cagaaagtta acaaaaattg atgacaagac tgctgactat 900
gttcgatctg gtctcactcc tcgatggagt gacctagatg ttaaccagca tgtgaataat 960
gtaaagtaca ttgggtggat cctggagagt gctccagtgg gaataatgga gaggcagaag 1020
ctgaaaagca tgactctgga gtatcggagg gaatgcggga gagacagtgt gcttcagtcc 1080
ctcactgcag ttacgggttg cgatatcggt aacctggcaa cagcggggga tgtggaatgt 1140
cagcatttgc tccgactcca ggatggagcg gaagtggtga gaggaagaac agagtggagt 1200
agtaaaacac caacaacaac ttggggaact gcaccgtaa 1239
<210> 34
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物 TEF-GSY-F
<400> 34
ctcgcaacaa ccgattccaa caagagaccg ggttggcggc gca 43
<210> 35
<211> 74
<212> DNA
<213> 人工序列
<220>
<223> 引物 TEF-GSY-R
<400> 35
ataacttcgt ataatgtatg ctatacgaag ttataagctt tgaatgattc ttatactcag 60
aaggaaatgc ttaa 74
<210> 36
<211> 1061
<212> DNA
<213> 莱茵衣藻
<400> 36
aagcttatgt ctgtcctcac caagtggctg ggtctcccct ctttcctgtt ctccgtcttc 60
gtgttctact ggtctctccc catcttcgcc attctgtacc gaatccgatt cgcttccctg 120
ggaaagcgaa acgacatgct cgactgggct cgagccctgg tcgcctactt ccgagtgacc 180
ctgctccagg ctggcgagca caccctgtac aagggcggtc cctgcctgta cctctgtaac 240
caccgatcct gggctgactt cttcattgac gcttacctga ccgagggacg agctgctctc 300
atgtctcgat ggctggtcta cttcgtgttc cccgtcttct gcacctcctg tatgatcctc 360
aagggtattg tcctgttcaa gcgaggaacc attgccgaca aggaagcctt caacgcctgg 420
ctggaccaga ccctgggatc ctctcacgtc cctggactgc tggtgtaccc cgagggacac 480
cgatctacca agcctgcctc cctgcctctc aagcgaggta tgctccacta cgctcactct 540
cgaaagctgc ccgtgcagat tgtcgtgacc cgaggcaagg acgaggtcct gtccgagaag 600
tctcagtccg tgcacttcgg acgaacctgc gtcaccacct tctctaaggt gctcaagtcc 660
gctgactacc ccaacttcga ggccttcttc accgacctgc aggctacctg ggactcttgt 720
tgggccgcta cctacggact ggaggacctc aagaacgtgc ctcgattctc tatgcccgga 780
cctcaggcct actcctactc ctcttccatg tgggtgcagc agctcgccat caccctcgtg 840
tctattctgg tcttcgctgg agtttgttac ggctcctggc gaggtctggc cgctgccctg 900
gctgctaccg gtgctgccca gcaggtggtt gctctggtgc tggctgcttg ggtgggttct 960
tccgtgctcc gatccttcct gtaacgatcg ttttttttta tatatatata tatatatata 1020
taactgtcta gaaataaaga gtatcatctt tcaaaaagct t 1061
<210> 37
<211> 935
<212> DNA
<213> 智人
<400> 37
aagcttatgg acctgtggcc cggagcttgg atgctgctcc tgctcctgtt cctcctcctg 60
ctgttcctcc tgcccaccct gtggttctgc tccccctctg ctaagtactt cttcaagatg 120
gccttctaca acggttggat tctgttcctg gccgtcctgg ctattcccgt ctgtgctgtg 180
cgaggacgaa acgtggagaa catgaagatc ctccgactga tgctcctgca catcaagtac 240
ctgtacggaa ttcgagttga ggtccgaggc gcccaccact tccctccctc ccagccttac 300
gtcgtggtct ctaaccacca gtcctctctg gacctcctgg gtatgatgga ggtgctccct 360
ggacgatgtg tccctatcgc taagcgagag ctgctctggg ctggttccgc tggactggct 420
tgttggctgg ctggcgtcat cttcattgac cgaaagcgaa ccggtgacgc tatttccgtg 480
atgtctgagg tggctcagac cctcctgacc caggacgttc gagtctgggt gttccctgag 540
ggaacccgaa accacaacgg ttccatgctg cccttcaagc gaggcgcctt ccacctcgct 600
gtccaggctc aggtccctat tgtgcccatt gtcatgtcct cttaccagga cttctactgc 660
aagaaggagc gacgattcac ctctggacag tgtcaggtcc gagtgctgcc tcccgtgcct 720
accgagggac tgacccccga cgacgttcct gctctggctg accgagtccg acactccatg 780
ctgaccgtgt tccgagagat ttctaccgac ggtcgaggcg gtggagacta cctcaagaag 840
cccggcggtg gaggctaacg atcgtttttt tttatatata tatatatata tatataactg 900
tctagaaata aagagtatca tctttcaaaa agctt 935
<210> 38
<211> 860
<212> DNA
<213> 拟南芥
<400> 38
aagcttatgt ccgacctgtc tggtgctgct acccccgagt ccacctaccc tgagcctgag 60
atcaagctgt cctctcgact ccgaggaatt tgcttctgtc tcgtcgccgg cgtgtctgct 120
atcgtcctga ttgtgctcat gatcaccggc caccccttcg tcctgctctt cgaccgatac 180
cgacgaaagt tccaccactt catcgccaag ctgtgggctt ccatctctat ctaccccttc 240
tacaagaccg acattcaggg tctggagaac ctcccctcct ctgacacccc ctgcgtctac 300
gtgtccaacc accagtcttt cctggacatc tacaccctgc tctccctcgg acagtcttac 360
aagttcattt ccaagaccgg catcttcgtc attcccgtga tcggctgggc catgtccatg 420
atgggtgtcg tgcccctgaa gcgaatggac ccccgatctc aggtcgactg cctgaagcga 480
tgtatggagc tcgtcaagaa gggtgcctcc gtcttcttct tccccgaggg aacccgatct 540
aaggacggac gactgggccc cttcaagaag ggcgctttca ccattgctgc taagaccggt 600
gtgcctgtgg tgcccattac cctgatgggc accggcaaga tcatgcccac cggttccgag 660
ggaattctca accacggtga cgtccgagtg atcattcaca agcccatcta cggatctaag 720
gctgacctgc tctgtgacga ggcccgaaac aagattgctg agtccatgaa cctgctctct 780
taacgatcgt ttttttttat atatatatat atatatatat aactgtctag aaataaagag 840
tatcatcttt caaaaagctt 860
<210> 39
<211> 852
<212> DNA
<213> 智人
<400> 39
atggatttgt ggccaggggc atggatgctg ctgctgctgc tcttcctgct gctgctcttc 60
ctgctgccca ccctgtggtt ctgcagcccc agtgccaagt acttcttcaa gatggccttc 120
tacaatggct ggatcctctt cctggctgtg ctcgccatcc ctgtgtgtgc cgtgcgagga 180
cgcaacgtcg agaacatgaa gatcttgcgt ctaatgctgc tccacatcaa atacctgtac 240
gggatccgag tggaggtgcg aggggctcac cacttccctc cctcgcagcc ctatgttgtt 300
gtctccaacc accagagctc tctcgatctg cttgggatga tggaggtact gccaggccgc 360
tgtgtgccca ttgccaagcg cgagctactg tgggctggct ctgccgggct ggcctgctgg 420
ctggcaggag tcatcttcat cgaccggaag cgcacggggg atgccatcag tgtcatgtct 480
gaggtcgccc agaccctgct cacccaggac gtgagggtct gggtgtttcc tgagggaacg 540
agaaaccaca atggctccat gctgcccttc aaacgtggcg ccttccatct tgcagtgcag 600
gcccaggttc ccattgtccc catagtcatg tcctcctacc aagacttcta ctgcaagaag 660
gagcgtcgct tcacctcggg acaatgtcag gtgcgggtgc tgcccccagt gcccacggaa 720
gggctgacac cagatgacgt cccagctctg gctgacagag tccggcactc catgctcact 780
gttttccggg aaatctccac tgatggccgg ggtggtggtg actatctgaa gaagcctggg 840
ggcggtgggt ga 852
<210> 40
<211> 283
<212> PRT
<213> 智人
<400> 40
Met Asp Leu Trp Pro Gly Ala Trp Met Leu Leu Leu Leu Leu Phe Leu
1 5 10 15
Leu Leu Leu Phe Leu Leu Pro Thr Leu Trp Phe Cys Ser Pro Ser Ala
20 25 30
Lys Tyr Phe Phe Lys Met Ala Phe Tyr Asn Gly Trp Ile Leu Phe Leu
35 40 45
Ala Val Leu Ala Ile Pro Val Cys Ala Val Arg Gly Arg Asn Val Glu
50 55 60
Asn Met Lys Ile Leu Arg Leu Met Leu Leu His Ile Lys Tyr Leu Tyr
65 70 75 80
Gly Ile Arg Val Glu Val Arg Gly Ala His His Phe Pro Pro Ser Gln
85 90 95
Pro Tyr Val Val Val Ser Asn His Gln Ser Ser Leu Asp Leu Leu Gly
100 105 110
Met Met Glu Val Leu Pro Gly Arg Cys Val Pro Ile Ala Lys Arg Glu
115 120 125
Leu Leu Trp Ala Gly Ser Ala Gly Leu Ala Cys Trp Leu Ala Gly Val
130 135 140
Ile Phe Ile Asp Arg Lys Arg Thr Gly Asp Ala Ile Ser Val Met Ser
145 150 155 160
Glu Val Ala Gln Thr Leu Leu Thr Gln Asp Val Arg Val Trp Val Phe
165 170 175
Pro Glu Gly Thr Arg Asn His Asn Gly Ser Met Leu Pro Phe Lys Arg
180 185 190
Gly Ala Phe His Leu Ala Val Gln Ala Gln Val Pro Ile Val Pro Ile
195 200 205
Val Met Ser Ser Tyr Gln Asp Phe Tyr Cys Lys Lys Glu Arg Arg Phe
210 215 220
Thr Ser Gly Gln Cys Gln Val Arg Val Leu Pro Pro Val Pro Thr Glu
225 230 235 240
Gly Leu Thr Pro Asp Asp Val Pro Ala Leu Ala Asp Arg Val Arg His
245 250 255
Ser Met Leu Thr Val Phe Arg Glu Ile Ser Thr Asp Gly Arg Gly Gly
260 265 270
Gly Asp Tyr Leu Lys Lys Pro Gly Gly Gly Gly
275 280
<210> 41
<211> 978
<212> DNA
<213> 莱茵衣藻
<400> 41
atgtccgtgc tcacaaagtg gctgggcctg ccctcattct tgttctccgt gtttgtgttc 60
tactggagtt tgcccatctt tgcgatccta tatcgtatcc gcttcgcctc tctggggaag 120
cgcaatgata tgctcgactg ggcccgcgcg ctcgtcgcct acttccgagt aacgctcctg 180
caggcggggg agcacacgct gtacaaggga gggccatgcc tgtacctgtg caaccaccgc 240
agctgggcgg acttcttcat tgacgcctac ctcactgagg gccgcgcggc gctcatgagc 300
aggtggctgg tgtatttcgt gttcccagtg ttctgtacct cgtgcatgat cctcaagggg 360
atagtgctgt tcaagcgcgg caccattgcg gataaagagg ccttcaacgc ctggcttgac 420
cagaccctgg gcagcagcca cgtgcctggg ctgctggtgt accccgaggg ccaccgcagc 480
accaagccgg ccagcctgcc gctgaagcgc ggcatgctgc actacgcgca cagccgcaag 540
ctgccggtgc agattgtcgt gactcgcggc aaggacgagg tgctgagtga gaagtcgcag 600
tcggtgcact tcggccgcac ctgcgtcacc accttctcca aggtgctcaa gtcggccgac 660
taccccaact ttgaagcctt cttcacggac ctgcaggcga cctgggactc gtgctgggca 720
gccacgtacg gcctggagga tctcaaaaac gtgccgcgct tctccatgcc ggggccccag 780
gcctacagct acagctccag catgtgggtc cagcagctcg ccatcacgct ggtcagcatc 840
ctagtctttg cgggcgtgtg ctacggctcc tggcgcggct tggcggcggc gctggcggcg 900
accggcgcgg cgcagcaggt ggtggcactg gtgctggcgg cctgggtggg cagctccgtg 960
cttcgtagct tcctctag 978
<210> 42
<211> 325
<212> PRT
<213> 莱茵衣藻
<400> 42
Met Ser Val Leu Thr Lys Trp Leu Gly Leu Pro Ser Phe Leu Phe Ser
1 5 10 15
Val Phe Val Phe Tyr Trp Ser Leu Pro Ile Phe Ala Ile Leu Tyr Arg
20 25 30
Ile Arg Phe Ala Ser Leu Gly Lys Arg Asn Asp Met Leu Asp Trp Ala
35 40 45
Arg Ala Leu Val Ala Tyr Phe Arg Val Thr Leu Leu Gln Ala Gly Glu
50 55 60
His Thr Leu Tyr Lys Gly Gly Pro Cys Leu Tyr Leu Cys Asn His Arg
65 70 75 80
Ser Trp Ala Asp Phe Phe Ile Asp Ala Tyr Leu Thr Glu Gly Arg Ala
85 90 95
Ala Leu Met Ser Arg Trp Leu Val Tyr Phe Val Phe Pro Val Phe Cys
100 105 110
Thr Ser Cys Met Ile Leu Lys Gly Ile Val Leu Phe Lys Arg Gly Thr
115 120 125
Ile Ala Asp Lys Glu Ala Phe Asn Ala Trp Leu Asp Gln Thr Leu Gly
130 135 140
Ser Ser His Val Pro Gly Leu Leu Val Tyr Pro Glu Gly His Arg Ser
145 150 155 160
Thr Lys Pro Ala Ser Leu Pro Leu Lys Arg Gly Met Leu His Tyr Ala
165 170 175
His Ser Arg Lys Leu Pro Val Gln Ile Val Val Thr Arg Gly Lys Asp
180 185 190
Glu Val Leu Ser Glu Lys Ser Gln Ser Val His Phe Gly Arg Thr Cys
195 200 205
Val Thr Thr Phe Ser Lys Val Leu Lys Ser Ala Asp Tyr Pro Asn Phe
210 215 220
Glu Ala Phe Phe Thr Asp Leu Gln Ala Thr Trp Asp Ser Cys Trp Ala
225 230 235 240
Ala Thr Tyr Gly Leu Glu Asp Leu Lys Asn Val Pro Arg Phe Ser Met
245 250 255
Pro Gly Pro Gln Ala Tyr Ser Tyr Ser Ser Ser Met Trp Val Gln Gln
260 265 270
Leu Ala Ile Thr Leu Val Ser Ile Leu Val Phe Ala Gly Val Cys Tyr
275 280 285
Gly Ser Trp Arg Gly Leu Ala Ala Ala Leu Ala Ala Thr Gly Ala Ala
290 295 300
Gln Gln Val Val Ala Leu Val Leu Ala Ala Trp Val Gly Ser Ser Val
305 310 315 320
Leu Arg Ser Phe Leu
325
<210> 43
<211> 1338
<212> DNA
<213> 微拟球藻
<400> 43
atgtctggtc cagccgatgc caggggccct gcccgagtat ccgataccat tattggtgcg 60
cccgaccgcc accatgtcgg tctgaccatc ctcgtcgccc ttctcctgag cgccttttgc 120
ttttgcgaat tcttcatgct gcccgtggcg atggcgttac tgcccgtgcc aggtacaaca 180
tcgaggaagt tgctgcggcg ctgggagggc ttggtagcca aaacttggct gtcctttgga 240
gggtggctcc tggaaaatgt gggtggggtc aagctcattc ttacgggcga ctcgatactc 300
cccggtgatc gtgtgctgat tctcagcaat cataggacga gaattgattg gatgtttcta 360
tggtgctggg ctgcacgttt cgacctgctt tcctcgtacc gagtcattct caagtcgtct 420
ttacgcactt tcccctggtg gggctggggg atgagtttat gtttgtttcc tttcatccac 480
cgaggaagca acaatcgcga cgcggatttg acccgtattg aacatatttg tcagtattta 540
agcgagctgg gtgtgccaaa ctccctcatt ctcttcccag aaggaactga cctttcccaa 600
tctaaccaaa aacgcgatcg tgattatgct ctagccaaaa acctacccat ttaccataac 660
gtgcttcacc cacgttcagg cgcctttatt gctagcttga cggccatgcg tcctcatttg 720
gatgctgttg tggatcttac tatcggctat gtcgattaca caccagggga aagaccttcc 780
gagctctctc ttctcaaggg ccggctaccg cacgaggtcc acataaatat gaaacgatgg 840
gacataaaga cgactccctt acttcgagag aacccatcgg accgggcgga acaatttttg 900
cgcgaatctt ttgatcggaa agaggcattg ctgacagcct tctacagcaa caacacggac 960
acttcaggtg gtgggctcaa gcaaacgtca tcattgctcg tatcctcttc ctcttcacct 1020
ttctccaata gcagtgcacc tttaggattt ccttccttct cctccccctc cacctccgtc 1080
gcctcttctt acaacgagcc agtgtatgta cacaatgctc actggacata cggcaaaggg 1140
cttttgggga cggggattgc catggcgttg gtgctgatgt tagcgtgctg cttgcctgct 1200
gttgcgtttt gcgggtatag tatcgttgtc tttttgttgt tcgtgacagt gaattcggtg 1260
tgggagggct tcgatacctt ggaattgaat ttggctgtgt ggtggggcgg gcaggcaggg 1320
caaacgaagc aagagtga 1338
<210> 44
<211> 445
<212> PRT
<213> 微拟球藻
<400> 44
Met Ser Gly Pro Ala Asp Ala Arg Gly Pro Ala Arg Val Ser Asp Thr
1 5 10 15
Ile Ile Gly Ala Pro Asp Arg His His Val Gly Leu Thr Ile Leu Val
20 25 30
Ala Leu Leu Leu Ser Ala Phe Cys Phe Cys Glu Phe Phe Met Leu Pro
35 40 45
Val Ala Met Ala Leu Leu Pro Val Pro Gly Thr Thr Ser Arg Lys Leu
50 55 60
Leu Arg Arg Trp Glu Gly Leu Val Ala Lys Thr Trp Leu Ser Phe Gly
65 70 75 80
Gly Trp Leu Leu Glu Asn Val Gly Gly Val Lys Leu Ile Leu Thr Gly
85 90 95
Asp Ser Ile Leu Pro Gly Asp Arg Val Leu Ile Leu Ser Asn His Arg
100 105 110
Thr Arg Ile Asp Trp Met Phe Leu Trp Cys Trp Ala Ala Arg Phe Asp
115 120 125
Leu Leu Ser Ser Tyr Arg Val Ile Leu Lys Ser Ser Leu Arg Thr Phe
130 135 140
Pro Trp Trp Gly Trp Gly Met Ser Leu Cys Leu Phe Pro Phe Ile His
145 150 155 160
Arg Gly Ser Asn Asn Arg Asp Ala Asp Leu Thr Arg Ile Glu His Ile
165 170 175
Cys Gln Tyr Leu Ser Glu Leu Gly Val Pro Asn Ser Leu Ile Leu Phe
180 185 190
Pro Glu Gly Thr Asp Leu Ser Gln Ser Asn Gln Lys Arg Asp Arg Asp
195 200 205
Tyr Ala Leu Ala Lys Asn Leu Pro Ile Tyr His Asn Val Leu His Pro
210 215 220
Arg Ser Gly Ala Phe Ile Ala Ser Leu Thr Ala Met Arg Pro His Leu
225 230 235 240
Asp Ala Val Val Asp Leu Thr Ile Gly Tyr Val Asp Tyr Thr Pro Gly
245 250 255
Glu Arg Pro Ser Glu Leu Ser Leu Leu Lys Gly Arg Leu Pro His Glu
260 265 270
Val His Ile Asn Met Lys Arg Trp Asp Ile Lys Thr Thr Pro Leu Leu
275 280 285
Arg Glu Asn Pro Ser Asp Arg Ala Glu Gln Phe Leu Arg Glu Ser Phe
290 295 300
Asp Arg Lys Glu Ala Leu Leu Thr Ala Phe Tyr Ser Asn Asn Thr Asp
305 310 315 320
Thr Ser Gly Gly Gly Leu Lys Gln Thr Ser Ser Leu Leu Val Ser Ser
325 330 335
Ser Ser Ser Pro Phe Ser Asn Ser Ser Ala Pro Leu Gly Phe Pro Ser
340 345 350
Phe Ser Ser Pro Ser Thr Ser Val Ala Ser Ser Tyr Asn Glu Pro Val
355 360 365
Tyr Val His Asn Ala His Trp Thr Tyr Gly Lys Gly Leu Leu Gly Thr
370 375 380
Gly Ile Ala Met Ala Leu Val Leu Met Leu Ala Cys Cys Leu Pro Ala
385 390 395 400
Val Ala Phe Cys Gly Tyr Ser Ile Val Val Phe Leu Leu Phe Val Thr
405 410 415
Val Asn Ser Val Trp Glu Gly Phe Asp Thr Leu Glu Leu Asn Leu Ala
420 425 430
Val Trp Trp Gly Gly Gln Ala Gly Gln Thr Lys Gln Glu
435 440 445
<210> 45
<211> 1449
<212> DNA
<213> 微拟球藻
<400> 45
atgtggggct tattatggaa acacatgacc ctacgactcc tcttcgtctc agtctgcttg 60
gtcacctcct ccatcggcgc caacatgatg gccctcgccc tcttcctcgt catccgaccc 120
ttttcccgct cgctgtatcg acggttggtg tcacagtgtg tggcatgcat gtggatcgat 180
gcactgtcct tgctcctacc gggaaccaac atccacattg ctgctgactc tgacatgccc 240
gacgggatca ctgccggcat cgtggtggcc aatcaccaat acgaaggtga ttggtggttc 300
atgctcatgg tcgcccgctt cctgggcctg catgggaacg tgaagataat agtcagggag 360
ggccttaaga agatccccct cttggggtgg ctggtccggc ttgtcgagta cccattcatc 420
tcctcctcct ggtcgctctc ccgcacgaac ctctttgggc tgctgcggag tttcaatgcg 480
gacgacttcc ccgtcctcct cttccagttc cccgagggcg atcgcatcga tgccaaagtc 540
cgtcagcaat cgcttgcctt tgcggccaag gagcaacgcc cgcacctcct ccacgtcctc 600
ctcccccgta ctaccggttt caacacctgc atcgaagcgc ttaggacttc ccatcccccc 660
gtctatgata tgacaattgc catccctggc accacgggcc agccttcctc ctcctcctcc 720
tcctcctcac cttctgctgc agctgctgcg gcgtctccat ctgctggtgc ggcggcggcg 780
gcggcggcgg cgagcaacac atcaaccaac ccttcttcca accctccctc gagcagcagc 840
accgatacta gtccagctgc tgctgctgct gctgctgctg atgccgctgc tgccgagagt 900
catgacgctt ctctctttca caccttcctc cgtttctgca acggcgaggg cccacgggac 960
gtgcatatca gactcaagcg ctactccctc aacgacgtcc tggccgaccc ccactggctc 1020
gacaccaaat gggccgagaa agaccgcgtc ttgacctact tttcccgcca cgcctgcttt 1080
cctgctccgc tccctcccca catggccaat ggtgggggca gcactcagac acacgggggg 1140
agggggggcc ccagagggcc ggcgggctct catgccgtgg tgggagcagc gggtaacgcc 1200
cagtatctcc ggtcgttcaa ttcccggaaa ttcaaagcag aaacgagttt tttggcgttg 1260
gcacgactgt tactcacgcc gttgtgcctt cccttgttgt tgctgatggc ctgtccgctc 1320
atgactctgt atgtatgcgt gagcaccgtc cgaaggtggt tgggggagga gattgtgttc 1380
tcgccaggag gacgggaggg agggagggag ggcgacgccg tttacgagtc cgttaattgg 1440
agcgtttga 1449
<210> 46
<211> 482
<212> PRT
<213> 微拟球藻
<400> 46
Met Trp Gly Leu Leu Trp Lys His Met Thr Leu Arg Leu Leu Phe Val
1 5 10 15
Ser Val Cys Leu Val Thr Ser Ser Ile Gly Ala Asn Met Met Ala Leu
20 25 30
Ala Leu Phe Leu Val Ile Arg Pro Phe Ser Arg Ser Leu Tyr Arg Arg
35 40 45
Leu Val Ser Gln Cys Val Ala Cys Met Trp Ile Asp Ala Leu Ser Leu
50 55 60
Leu Leu Pro Gly Thr Asn Ile His Ile Ala Ala Asp Ser Asp Met Pro
65 70 75 80
Asp Gly Ile Thr Ala Gly Ile Val Val Ala Asn His Gln Tyr Glu Gly
85 90 95
Asp Trp Trp Phe Met Leu Met Val Ala Arg Phe Leu Gly Leu His Gly
100 105 110
Asn Val Lys Ile Ile Val Arg Glu Gly Leu Lys Lys Ile Pro Leu Leu
115 120 125
Gly Trp Leu Val Arg Leu Val Glu Tyr Pro Phe Ile Ser Ser Ser Trp
130 135 140
Ser Leu Ser Arg Thr Asn Leu Phe Gly Leu Leu Arg Ser Phe Asn Ala
145 150 155 160
Asp Asp Phe Pro Val Leu Leu Phe Gln Phe Pro Glu Gly Asp Arg Ile
165 170 175
Asp Ala Lys Val Arg Gln Gln Ser Leu Ala Phe Ala Ala Lys Glu Gln
180 185 190
Arg Pro His Leu Leu His Val Leu Leu Pro Arg Thr Thr Gly Phe Asn
195 200 205
Thr Cys Ile Glu Ala Leu Arg Thr Ser His Pro Pro Val Tyr Asp Met
210 215 220
Thr Ile Ala Ile Pro Gly Thr Thr Gly Gln Pro Ser Ser Ser Ser Ser
225 230 235 240
Ser Ser Ser Pro Ser Ala Ala Ala Ala Ala Ala Ser Pro Ser Ala Gly
245 250 255
Ala Ala Ala Ala Ala Ala Ala Ala Ser Asn Thr Ser Thr Asn Pro Ser
260 265 270
Ser Asn Pro Pro Ser Ser Ser Ser Thr Asp Thr Ser Pro Ala Ala Ala
275 280 285
Ala Ala Ala Ala Ala Asp Ala Ala Ala Ala Glu Ser His Asp Ala Ser
290 295 300
Leu Phe His Thr Phe Leu Arg Phe Cys Asn Gly Glu Gly Pro Arg Asp
305 310 315 320
Val His Ile Arg Leu Lys Arg Tyr Ser Leu Asn Asp Val Leu Ala Asp
325 330 335
Pro His Trp Leu Asp Thr Lys Trp Ala Glu Lys Asp Arg Val Leu Thr
340 345 350
Tyr Phe Ser Arg His Ala Cys Phe Pro Ala Pro Leu Pro Pro His Met
355 360 365
Ala Asn Gly Gly Gly Ser Thr Gln Thr His Gly Gly Arg Gly Gly Pro
370 375 380
Arg Gly Pro Ala Gly Ser His Ala Val Val Gly Ala Ala Gly Asn Ala
385 390 395 400
Gln Tyr Leu Arg Ser Phe Asn Ser Arg Lys Phe Lys Ala Glu Thr Ser
405 410 415
Phe Leu Ala Leu Ala Arg Leu Leu Leu Thr Pro Leu Cys Leu Pro Leu
420 425 430
Leu Leu Leu Met Ala Cys Pro Leu Met Thr Leu Tyr Val Cys Val Ser
435 440 445
Thr Val Arg Arg Trp Leu Gly Glu Glu Ile Val Phe Ser Pro Gly Gly
450 455 460
Arg Glu Gly Gly Arg Glu Gly Asp Ala Val Tyr Glu Ser Val Asn Trp
465 470 475 480
Ser Val
<210> 47
<211> 1911
<212> DNA
<213> 微拟球藻
<400> 47
atgccgacct atgcggaccc tcccaggtcg cgcgcctacc agcacgaaag ctcctcctac 60
agcattgata atccgatccg cagcacccac gcctccagcc cacccgacta tttcaatcaa 120
aactccctcc gtgagtcccc ctccctccac tcctcatcct cagaatccga gatggactct 180
cagagcaatg gcaatggcag gacccctcaa cagcggcgag gcgacattca aaggcagaca 240
aacaaccctt ccgcctccgg cacgaacggc accccccgag gagcattcaa gagcacaatc 300
ttctccatgg cccacgctgt cgtcctctgc tggctcgtga tggccatgtc cttcggcatc 360
aatattttga atttcctcgt ttttgtaacc gtccgtcctt ggtcccgcct gacggcccga 420
aggctcatcg gtcactggtg gcaacggatg tgggtctcag taatgagcta cttgctccct 480
aagagcgaga tgattctgac aggggacatc gtgcttaaca ctgtggattc ctctcgcccg 540
gccatcatca tcgcgaatca ccaggtggat gcggattttt ggtatgtctg ggaagtggcg 600
cgggcctacg gaatgcatgg acggttgaag atcattttga aagccgattt ggctgtggtc 660
cccatcgtgg gctgggggat gaaacaattt gaattttgtt ttttgcaacg caattggcag 720
agggaccgga gggcgttcac gcgcttgttg tcgtcgtttg tcgaggacgg gtataagtgt 780
gccttgctcc tctttccaga gggcacgacc atcaacactg aagctgtcac caagtcccac 840
cgcttcggcc gtgagcagaa gaggcctcac ctcgaccatt gcatcctacc gcgttccacg 900
ggctttgcgg ctttggtcga gacaatgtgc cagtcccccc atggccattc tcctgttatt 960
tacgatctga ccgtcgccta tcacggttac tcaggcgaag ttcccaccta tgagatgggc 1020
tacgaccgag aacaagacat ggacgtcccg aacgtgttca aaatgataca gggacgggca 1080
tgtccaaagg ttcatgtcca cgtcaagacg tacgaagtcg atgaggcctt ggttaataat 1140
cccgagcagt ggttggatgc tcgatggttg gaaaaagacg cgctgctgga acggtttatt 1200
cagacgcaga gttttagggg ggagaaggag ggagggacac ggatcattca tcctcagggg 1260
agcttggcct cgctgctttt tttgttgatg ttgccgctga ttatcaccgt cctccttcct 1320
gccctgctct tgatcacttt ggtggcctgg cctgtgattc ttctggcggg cacgctcaac 1380
tttctgaact ttgtctcgcg ttcggccatg ggtgtggtag tcgggagcag cagcagccgt 1440
gggtctttct ccacgtcttg gcatcagtcc tcccaacacc ggcaccagca ccagcaacgg 1500
gtcagtggca atagcaccag tagcaccagt accagcaaca gtaccagcag tagcattggt 1560
cggaaggaag gggcattgaa ctgcaggcga agttctcgcc gaagccccag ccgtgaggga 1620
tgtggccacg ccaatggaga aggaccgagc tgtcctctcc tctctgtaag cctgccgcct 1680
ctccctgccg cagctgctgc tgccgctcct gctgctcctg cttcttcggc tccatcttct 1740
ccgctattgg ggagcacgcc gtcgccacgc cggtccccac ggtggaagac tccggcgggg 1800
atatcccctg tagtgcagag ctacgggaac aggaagggga aggaaggagg atggggtcta 1860
atggctgcgg cggccgcagc ggcaacgacg gaagaggagg atcggtacta g 1911
<210> 48
<211> 636
<212> PRT
<213> 微拟球藻
<400> 48
Met Pro Thr Tyr Ala Asp Pro Pro Arg Ser Arg Ala Tyr Gln His Glu
1 5 10 15
Ser Ser Ser Tyr Ser Ile Asp Asn Pro Ile Arg Ser Thr His Ala Ser
20 25 30
Ser Pro Pro Asp Tyr Phe Asn Gln Asn Ser Leu Arg Glu Ser Pro Ser
35 40 45
Leu His Ser Ser Ser Ser Glu Ser Glu Met Asp Ser Gln Ser Asn Gly
50 55 60
Asn Gly Arg Thr Pro Gln Gln Arg Arg Gly Asp Ile Gln Arg Gln Thr
65 70 75 80
Asn Asn Pro Ser Ala Ser Gly Thr Asn Gly Thr Pro Arg Gly Ala Phe
85 90 95
Lys Ser Thr Ile Phe Ser Met Ala His Ala Val Val Leu Cys Trp Leu
100 105 110
Val Met Ala Met Ser Phe Gly Ile Asn Ile Leu Asn Phe Leu Val Phe
115 120 125
Val Thr Val Arg Pro Trp Ser Arg Leu Thr Ala Arg Arg Leu Ile Gly
130 135 140
His Trp Trp Gln Arg Met Trp Val Ser Val Met Ser Tyr Leu Leu Pro
145 150 155 160
Lys Ser Glu Met Ile Leu Thr Gly Asp Ile Val Leu Asn Thr Val Asp
165 170 175
Ser Ser Arg Pro Ala Ile Ile Ile Ala Asn His Gln Val Asp Ala Asp
180 185 190
Phe Trp Tyr Val Trp Glu Val Ala Arg Ala Tyr Gly Met His Gly Arg
195 200 205
Leu Lys Ile Ile Leu Lys Ala Asp Leu Ala Val Val Pro Ile Val Gly
210 215 220
Trp Gly Met Lys Gln Phe Glu Phe Cys Phe Leu Gln Arg Asn Trp Gln
225 230 235 240
Arg Asp Arg Arg Ala Phe Thr Arg Leu Leu Ser Ser Phe Val Glu Asp
245 250 255
Gly Tyr Lys Cys Ala Leu Leu Leu Phe Pro Glu Gly Thr Thr Ile Asn
260 265 270
Thr Glu Ala Val Thr Lys Ser His Arg Phe Gly Arg Glu Gln Lys Arg
275 280 285
Pro His Leu Asp His Cys Ile Leu Pro Arg Ser Thr Gly Phe Ala Ala
290 295 300
Leu Val Glu Thr Met Cys Gln Ser Pro His Gly His Ser Pro Val Ile
305 310 315 320
Tyr Asp Leu Thr Val Ala Tyr His Gly Tyr Ser Gly Glu Val Pro Thr
325 330 335
Tyr Glu Met Gly Tyr Asp Arg Glu Gln Asp Met Asp Val Pro Asn Val
340 345 350
Phe Lys Met Ile Gln Gly Arg Ala Cys Pro Lys Val His Val His Val
355 360 365
Lys Thr Tyr Glu Val Asp Glu Ala Leu Val Asn Asn Pro Glu Gln Trp
370 375 380
Leu Asp Ala Arg Trp Leu Glu Lys Asp Ala Leu Leu Glu Arg Phe Ile
385 390 395 400
Gln Thr Gln Ser Phe Arg Gly Glu Lys Glu Gly Gly Thr Arg Ile Ile
405 410 415
His Pro Gln Gly Ser Leu Ala Ser Leu Leu Phe Leu Leu Met Leu Pro
420 425 430
Leu Ile Ile Thr Val Leu Leu Pro Ala Leu Leu Leu Ile Thr Leu Val
435 440 445
Ala Trp Pro Val Ile Leu Leu Ala Gly Thr Leu Asn Phe Leu Asn Phe
450 455 460
Val Ser Arg Ser Ala Met Gly Val Val Val Gly Ser Ser Ser Ser Arg
465 470 475 480
Gly Ser Phe Ser Thr Ser Trp His Gln Ser Ser Gln His Arg His Gln
485 490 495
His Gln Gln Arg Val Ser Gly Asn Ser Thr Ser Ser Thr Ser Thr Ser
500 505 510
Asn Ser Thr Ser Ser Ser Ile Gly Arg Lys Glu Gly Ala Leu Asn Cys
515 520 525
Arg Arg Ser Ser Arg Arg Ser Pro Ser Arg Glu Gly Cys Gly His Ala
530 535 540
Asn Gly Glu Gly Pro Ser Cys Pro Leu Leu Ser Val Ser Leu Pro Pro
545 550 555 560
Leu Pro Ala Ala Ala Ala Ala Ala Ala Pro Ala Ala Pro Ala Ser Ser
565 570 575
Ala Pro Ser Ser Pro Leu Leu Gly Ser Thr Pro Ser Pro Arg Arg Ser
580 585 590
Pro Arg Trp Lys Thr Pro Ala Gly Ile Ser Pro Val Val Gln Ser Tyr
595 600 605
Gly Asn Arg Lys Gly Lys Glu Gly Gly Trp Gly Leu Met Ala Ala Ala
610 615 620
Ala Ala Ala Ala Thr Thr Glu Glu Glu Asp Arg Tyr
625 630 635
<210> 49
<211> 678
<212> DNA
<213> 集胞藻属 PCC6803
<400> 49
gtggattccg agattaatca tcgtggtggt ttgagtgctc cccgcccaag ggaaacgtca 60
cttaatttag ctctctaccg gggcttgaaa tggggggtgg tgcggccact gctccatgga 120
ttgttccagg cccaggtata tggtcaggaa ttggtgccaa cccgggggcc ggccttggtg 180
gtgagcaacc atgccagtta ttttgacccc ccatttttgt cctgtgccat ggcccggccg 240
gtggccttta tggccaagga agagttattt aatgtgcccc tgctgggtcc agccattcgc 300
ctctatgggg cctatccagt caaacggggc agtggcgatc ggggagcatt gcgggccgcc 360
ttgacggcgc tgggggatgg ttggttagtg ggggtctttc tggagggaac cagaacaaag 420
gatggccgca ttcaccagcc aaaattgggg gctgccatga ttgcagctaa agcccaagtg 480
cccattattc ccgtcagcct agggggagta gagcaaattt ttcagcccgg ttccccctgg 540
ccccatcctg tgcctttaac tattcgcatt ggtaaggcga tcgcccctcc agtaaagaat 600
aggaaacccg aattggaagc ggttactaaa gcttgccaag cccaaattca cgagatgctg 660
gatttaggca gggattag 678
<210> 50
<211> 225
<212> PRT
<213> 集胞藻属 PCC6803
<400> 50
Met Asp Ser Glu Ile Asn His Arg Gly Gly Leu Ser Ala Pro Arg Pro
1 5 10 15
Arg Glu Thr Ser Leu Asn Leu Ala Leu Tyr Arg Gly Leu Lys Trp Gly
20 25 30
Val Val Arg Pro Leu Leu His Gly Leu Phe Gln Ala Gln Val Tyr Gly
35 40 45
Gln Glu Leu Val Pro Thr Arg Gly Pro Ala Leu Val Val Ser Asn His
50 55 60
Ala Ser Tyr Phe Asp Pro Pro Phe Leu Ser Cys Ala Met Ala Arg Pro
65 70 75 80
Val Ala Phe Met Ala Lys Glu Glu Leu Phe Asn Val Pro Leu Leu Gly
85 90 95
Pro Ala Ile Arg Leu Tyr Gly Ala Tyr Pro Val Lys Arg Gly Ser Gly
100 105 110
Asp Arg Gly Ala Leu Arg Ala Ala Leu Thr Ala Leu Gly Asp Gly Trp
115 120 125
Leu Val Gly Val Phe Leu Glu Gly Thr Arg Thr Lys Asp Gly Arg Ile
130 135 140
His Gln Pro Lys Leu Gly Ala Ala Met Ile Ala Ala Lys Ala Gln Val
145 150 155 160
Pro Ile Ile Pro Val Ser Leu Gly Gly Val Glu Gln Ile Phe Gln Pro
165 170 175
Gly Ser Pro Trp Pro His Pro Val Pro Leu Thr Ile Arg Ile Gly Lys
180 185 190
Ala Ile Ala Pro Pro Val Lys Asn Arg Lys Pro Glu Leu Glu Ala Val
195 200 205
Thr Lys Ala Cys Gln Ala Gln Ile His Glu Met Leu Asp Leu Gly Arg
210 215 220
Asp
225
<210> 51
<211> 1170
<212> DNA
<213> 拟南芥
<400> 51
atggtgattg ctgcagctgt catcgtgcct ttgggccttc tcttcttcat atctggtctc 60
gctgtcaatc tctttcaggc agtttgctat gtactcattc gaccactgtc taagaacaca 120
tacagaaaaa ttaaccgggt ggttgcagaa accttgtggt tggagcttgt atggatagtt 180
gactggtggg ctggagttaa gatccaagtg tttgctgata atgagacctt caatcgaatg 240
ggcaaagaac atgctcttgt cgtttgtaat caccgaagtg atattgattg gcttgtggga 300
tggattctgg ctcagcggtc aggttgcctg ggaagcgcat tagctgtaat gaagaagtct 360
tccaaattcc ttccagtcat aggctggtca atgtggttct cggagtatct ctttctggaa 420
agaaattggg ccaaggatga aagcactcta aagtcaggtc ttcagcgctt gagcgacttc 480
cctcgacctt tctggttagc cctttttgtg gagggaactc gctttacaga agccaaactt 540
aaagccgcac aagagtatgc agcctcctct gaattgccta tccctcgaaa tgtgttgatt 600
cctcgcacca aaggtttcgt gtcagctgtt agtaatatgc gttcatttgt cccagcaatt 660
tatgatatga cagtgactat tccaaaaacc tctccaccac ccacgatgct aagactattc 720
aaaggacaac cttcagtggt gcatgttcac atcaagtgtc actcgatgaa agacttacct 780
gaatcagatg acgcaattgc acagtggtgc agagatcagt ttgtggctaa ggatgctctg 840
ttagacaaac acatagctgc agacactttc cccggtcaac aagaacagaa cattggccgt 900
cccataaagt cccttgcggt ggttctatca tgggcatgcg tactaactct tggagcaata 960
aagttcctac actgggcaca actcttttct tcatggaaag gtatcacgat atcggcgctt 1020
ggtctaggta tcatcactct ctgtatgcag atcctgatac gctcgtctca gtcagagcgt 1080
tcgaccccag ccaaagtcgt cccagccaag ccaaaagaca atcaccaccc agaatcatcc 1140
tcccaaacag aaacggagaa ggagaagtaa 1170
<210> 52
<211> 389
<212> PRT
<213> 拟南芥
<400> 52
Met Val Ile Ala Ala Ala Val Ile Val Pro Leu Gly Leu Leu Phe Phe
1 5 10 15
Ile Ser Gly Leu Ala Val Asn Leu Phe Gln Ala Val Cys Tyr Val Leu
20 25 30
Ile Arg Pro Leu Ser Lys Asn Thr Tyr Arg Lys Ile Asn Arg Val Val
35 40 45
Ala Glu Thr Leu Trp Leu Glu Leu Val Trp Ile Val Asp Trp Trp Ala
50 55 60
Gly Val Lys Ile Gln Val Phe Ala Asp Asn Glu Thr Phe Asn Arg Met
65 70 75 80
Gly Lys Glu His Ala Leu Val Val Cys Asn His Arg Ser Asp Ile Asp
85 90 95
Trp Leu Val Gly Trp Ile Leu Ala Gln Arg Ser Gly Cys Leu Gly Ser
100 105 110
Ala Leu Ala Val Met Lys Lys Ser Ser Lys Phe Leu Pro Val Ile Gly
115 120 125
Trp Ser Met Trp Phe Ser Glu Tyr Leu Phe Leu Glu Arg Asn Trp Ala
130 135 140
Lys Asp Glu Ser Thr Leu Lys Ser Gly Leu Gln Arg Leu Ser Asp Phe
145 150 155 160
Pro Arg Pro Phe Trp Leu Ala Leu Phe Val Glu Gly Thr Arg Phe Thr
165 170 175
Glu Ala Lys Leu Lys Ala Ala Gln Glu Tyr Ala Ala Ser Ser Glu Leu
180 185 190
Pro Ile Pro Arg Asn Val Leu Ile Pro Arg Thr Lys Gly Phe Val Ser
195 200 205
Ala Val Ser Asn Met Arg Ser Phe Val Pro Ala Ile Tyr Asp Met Thr
210 215 220
Val Thr Ile Pro Lys Thr Ser Pro Pro Pro Thr Met Leu Arg Leu Phe
225 230 235 240
Lys Gly Gln Pro Ser Val Val His Val His Ile Lys Cys His Ser Met
245 250 255
Lys Asp Leu Pro Glu Ser Asp Asp Ala Ile Ala Gln Trp Cys Arg Asp
260 265 270
Gln Phe Val Ala Lys Asp Ala Leu Leu Asp Lys His Ile Ala Ala Asp
275 280 285
Thr Phe Pro Gly Gln Gln Glu Gln Asn Ile Gly Arg Pro Ile Lys Ser
290 295 300
Leu Ala Val Val Leu Ser Trp Ala Cys Val Leu Thr Leu Gly Ala Ile
305 310 315 320
Lys Phe Leu His Trp Ala Gln Leu Phe Ser Ser Trp Lys Gly Ile Thr
325 330 335
Ile Ser Ala Leu Gly Leu Gly Ile Ile Thr Leu Cys Met Gln Ile Leu
340 345 350
Ile Arg Ser Ser Gln Ser Glu Arg Ser Thr Pro Ala Lys Val Val Pro
355 360 365
Ala Lys Pro Lys Asp Asn His His Pro Glu Ser Ser Ser Gln Thr Glu
370 375 380
Thr Glu Lys Glu Lys
385
<210> 53
<211> 1149
<212> DNA
<213> 向日葵
<400> 53
atggctatcg cagcagcagc tgttatcgtc cctattggcg tcctcttctt cgtctccggc 60
ctcatcgtca atctcattca ggcgattata tttgtgactg tacgaccgtt ctcgaagagc 120
ttgtttaggc ggattaacag acaggtagct gagttgttgt ggctggagct tgtgtggatt 180
gttgattggt gggctggagt taaggttaac ctgtacacag atgccgagac cctgaagatg 240
atgggtaaag aacatgctct tgtgatagct aatcataaaa gtgacattga ttggctcatt 300
ggatgggtgt ttgctcagag atcaggttgt cttggtagca cattggctgt catgaagaaa 360
tcatcgaagt ttcttcccgt cattggttgg tcaatgtggt tttctgagta tctttttctt 420
gagagaagtt gggctaaaga tgaaagtacc ttgaagtcag gcctccgacg tctaaaagat 480
taccctcaac ccttttggtt ggcccttttt gttgaaggga ctcgctttac taaagcaaaa 540
cttttagcag ctcaagaata tgcatcttca atgggattac ctgttcccag aaatgtctta 600
attccaagaa caaagggatt tgttacttca gtgagtgaaa tgagatcatt tgctcctgca 660
atttacgata tgacggttgc gattcccaaa gattcaactc cgccaacaat gctgcgcctc 720
tttaaagggc agtcgtctgt gattcacgtt aaagttaagc gtcatttaat gaaggacctg 780
ccagaaacag atgaaggtgt tgcacaatgg tgtaaagata tttttgttgc caaggatgat 840
atattagata aacataaaga attaaacgcc tttcctgatt cagaactcca tgaaattggc 900
cgaccattga agtctcttgt ggtggttgta tcttgggcat gtctgcttgt actcgggatc 960
ttcaagttcc tgcaatggtc taatctttta tcctcatgga aggggctcac attcactgca 1020
attgggttgg ggattgttac ctttttaatg caaatcttga ttcagttttc gcaatctgaa 1080
cgttctacac ctgcaaaagt ggcccccaca aggtctagca atggtaatgt acaagagaaa 1140
ctgcactga 1149
<210> 54
<211> 382
<212> PRT
<213> 向日葵
<400> 54
Met Ala Ile Ala Ala Ala Ala Val Ile Val Pro Ile Gly Val Leu Phe
1 5 10 15
Phe Val Ser Gly Leu Ile Val Asn Leu Ile Gln Ala Ile Ile Phe Val
20 25 30
Thr Val Arg Pro Phe Ser Lys Ser Leu Phe Arg Arg Ile Asn Arg Gln
35 40 45
Val Ala Glu Leu Leu Trp Leu Glu Leu Val Trp Ile Val Asp Trp Trp
50 55 60
Ala Gly Val Lys Val Asn Leu Tyr Thr Asp Ala Glu Thr Leu Lys Met
65 70 75 80
Met Gly Lys Glu His Ala Leu Val Ile Ala Asn His Lys Ser Asp Ile
85 90 95
Asp Trp Leu Ile Gly Trp Val Phe Ala Gln Arg Ser Gly Cys Leu Gly
100 105 110
Ser Thr Leu Ala Val Met Lys Lys Ser Ser Lys Phe Leu Pro Val Ile
115 120 125
Gly Trp Ser Met Trp Phe Ser Glu Tyr Leu Phe Leu Glu Arg Ser Trp
130 135 140
Ala Lys Asp Glu Ser Thr Leu Lys Ser Gly Leu Arg Arg Leu Lys Asp
145 150 155 160
Tyr Pro Gln Pro Phe Trp Leu Ala Leu Phe Val Glu Gly Thr Arg Phe
165 170 175
Thr Lys Ala Lys Leu Leu Ala Ala Gln Glu Tyr Ala Ser Ser Met Gly
180 185 190
Leu Pro Val Pro Arg Asn Val Leu Ile Pro Arg Thr Lys Gly Phe Val
195 200 205
Thr Ser Val Ser Glu Met Arg Ser Phe Ala Pro Ala Ile Tyr Asp Met
210 215 220
Thr Val Ala Ile Pro Lys Asp Ser Thr Pro Pro Thr Met Leu Arg Leu
225 230 235 240
Phe Lys Gly Gln Ser Ser Val Ile His Val Lys Val Lys Arg His Leu
245 250 255
Met Lys Asp Leu Pro Glu Thr Asp Glu Gly Val Ala Gln Trp Cys Lys
260 265 270
Asp Ile Phe Val Ala Lys Asp Asp Ile Leu Asp Lys His Lys Glu Leu
275 280 285
Asn Ala Phe Pro Asp Ser Glu Leu His Glu Ile Gly Arg Pro Leu Lys
290 295 300
Ser Leu Val Val Val Val Ser Trp Ala Cys Leu Leu Val Leu Gly Ile
305 310 315 320
Phe Lys Phe Leu Gln Trp Ser Asn Leu Leu Ser Ser Trp Lys Gly Leu
325 330 335
Thr Phe Thr Ala Ile Gly Leu Gly Ile Val Thr Phe Leu Met Gln Ile
340 345 350
Leu Ile Gln Phe Ser Gln Ser Glu Arg Ser Thr Pro Ala Lys Val Ala
355 360 365
Pro Thr Arg Ser Ser Asn Gly Asn Val Gln Glu Lys Leu His
370 375 380
<210> 55
<211> 1517
<212> DNA
<213> 亚麻荠
<400> 55
ctccttataa tttcaatcgc tcccatttaa tcgcaaagtt atttttattt tatttttctg 60
gttggcttat tttttatttt attttatgtt ttaaatgttc tgcaacttgt cttgaaatct 120
ggggtaaaaa gaaaaaagaa aaaaaaaact tcgctattgg tttttctgga aatctcagaa 180
acgattattt tgggttttct atacttcgct tcctcacatt agagcttcgg tgttcttttt 240
tttccttttc tttttctttt tatttgggtg tgagggacat ttttccatgg tgattgctgc 300
agctgtcatc gtgcctttgg gccttctctt cttcatatct ggtctcgttg tcaatctcat 360
tcaggcactt tgctatgtcc tcattcggcc actgtctaag aacacttacc ggaaaatcaa 420
ccgggtggtt gctgaaacct tgtggttgga gcttgtttgg attgttgatt ggtgggctgg 480
agtaaagatc caagtgtttg ctgataatga gaccttcaat cgaatgggca aagaacacgc 540
tcttgtcgtt tgtaatcacc gtagtgatat tgattggctt gttggatggg ttctggctca 600
gcggtcaggt tccctgggaa gcgctttggc tgtaatgaag aagtcttcca aattccttcc 660
agtcataggc tggtcaatgt ggttctcaga gtatctgttt ctggaaagaa attggtccaa 720
ggatgaaagc actctaaagt caggtcttca gcgcttgagt gactttcctc gacctttctg 780
gctagccctt tttgtggagg gaactcgctt tacagaggct aaactcaaag cagcacaaga 840
gtatgcagcc tcctctgact tgcctatccc tcgaaatgtg ttgattcctc gcaccaaagg 900
ttttgtgtca gctgttagta atatgcgttc atttgtccca gccatttatg atatgacagt 960
gactattcca aaaacttctc caccgcccac gatgctaaga ctattcaaag gacaaccttc 1020
tgtggtacat gttcacatca agtgtcactc catgaaagac ttgcctgaat cagatgacgc 1080
aattgcacag tggtgcagag atcagtttgt ggctaaggat gctttgttag acaaacacat 1140
agctgcagac actttccccg gtcaacagga acagaacatt ggccgtccca taaagtccct 1200
tgcggtggtt ctatcatggg catgcgtatt aactcttgga gcaattaagt tcctacactg 1260
ggcacaactc ttttcgtcat ggaaaggtat cgcgctatcg gggcttgctc tgggtatcat 1320
cactctcggt atgcagatcc tgatacgctc gtctcagtca gagcgttcaa ccccagccaa 1380
agtggttcca gcaaagccaa aggaccatca caactcagaa tcatcctccc aaacagaagt 1440
ggagaagcag aagtaaaaaa agtggatatc aaagatcaaa caacaaacag aagaagaaaa 1500
gcgtatcagt tttgtta 1517
<210> 56
<211> 389
<212> PRT
<213> 亚麻荠
<400> 56
Met Val Ile Ala Ala Ala Val Ile Val Pro Leu Gly Leu Leu Phe Phe
1 5 10 15
Ile Ser Gly Leu Val Val Asn Leu Ile Gln Ala Leu Cys Tyr Val Leu
20 25 30
Ile Arg Pro Leu Ser Lys Asn Thr Tyr Arg Lys Ile Asn Arg Val Val
35 40 45
Ala Glu Thr Leu Trp Leu Glu Leu Val Trp Ile Val Asp Trp Trp Ala
50 55 60
Gly Val Lys Ile Gln Val Phe Ala Asp Asn Glu Thr Phe Asn Arg Met
65 70 75 80
Gly Lys Glu His Ala Leu Val Val Cys Asn His Arg Ser Asp Ile Asp
85 90 95
Trp Leu Val Gly Trp Val Leu Ala Gln Arg Ser Gly Ser Leu Gly Ser
100 105 110
Ala Leu Ala Val Met Lys Lys Ser Ser Lys Phe Leu Pro Val Ile Gly
115 120 125
Trp Ser Met Trp Phe Ser Glu Tyr Leu Phe Leu Glu Arg Asn Trp Ser
130 135 140
Lys Asp Glu Ser Thr Leu Lys Ser Gly Leu Gln Arg Leu Ser Asp Phe
145 150 155 160
Pro Arg Pro Phe Trp Leu Ala Leu Phe Val Glu Gly Thr Arg Phe Thr
165 170 175
Glu Ala Lys Leu Lys Ala Ala Gln Glu Tyr Ala Ala Ser Ser Asp Leu
180 185 190
Pro Ile Pro Arg Asn Val Leu Ile Pro Arg Thr Lys Gly Phe Val Ser
195 200 205
Ala Val Ser Asn Met Arg Ser Phe Val Pro Ala Ile Tyr Asp Met Thr
210 215 220
Val Thr Ile Pro Lys Thr Ser Pro Pro Pro Thr Met Leu Arg Leu Phe
225 230 235 240
Lys Gly Gln Pro Ser Val Val His Val His Ile Lys Cys His Ser Met
245 250 255
Lys Asp Leu Pro Glu Ser Asp Asp Ala Ile Ala Gln Trp Cys Arg Asp
260 265 270
Gln Phe Val Ala Lys Asp Ala Leu Leu Asp Lys His Ile Ala Ala Asp
275 280 285
Thr Phe Pro Gly Gln Gln Glu Gln Asn Ile Gly Arg Pro Ile Lys Ser
290 295 300
Leu Ala Val Val Leu Ser Trp Ala Cys Val Leu Thr Leu Gly Ala Ile
305 310 315 320
Lys Phe Leu His Trp Ala Gln Leu Phe Ser Ser Trp Lys Gly Ile Ala
325 330 335
Leu Ser Gly Leu Ala Leu Gly Ile Ile Thr Leu Gly Met Gln Ile Leu
340 345 350
Ile Arg Ser Ser Gln Ser Glu Arg Ser Thr Pro Ala Lys Val Val Pro
355 360 365
Ala Lys Pro Lys Asp His His Asn Ser Glu Ser Ser Ser Gln Thr Glu
370 375 380
Val Glu Lys Gln Lys
385
<210> 57
<211> 1173
<212> DNA
<213> 欧洲油菜
<400> 57
atggcgatgg cagcagcagt gattgtgcct ttggggattc tcttcttcat ttctggcctc 60
gttgtcaatc tcctccaggc agtttgctat gtcctcgttc gacctatgtc taagaacaca 120
tacagaaaga tcaaccgggt ggttgcagaa accttgtggt tggagcttgt ctggatcgtt 180
gactggtggg ctggagtcaa gatccaagtc tttgctgatg atgagacctt taatcgaatg 240
ggcaaagaac atgctcttgt cgtttgtaat caccgaagtg atattgattg gcttgttgga 300
tggattctcg ctcagaggtc aggttgcctg ggaagcgcat tagctgtaat gaagaagtct 360
tccaaatttc tcccagtcat aggctggtca atgtggttct ccgagtatct gtttcttgaa 420
agaaattggg caaaggatga aagcacttta cagtcaggtc ttcaacgctt gaacgacttc 480
ccacggcctt tctggctagc tctttttgtg gagggaaccc gcttcacaga ggcaaaactt 540
aaagcagcac aagagtacgc agcctcctct gagttgcctg tccctcgaaa tgtgttgatt 600
cctcgcacca aaggttttgt gtcagctgtt agtaacatgc gttcatttgt gccagccata 660
tatgatatga ccgtggctat tccaaaaact tctccacccc caacgatgct aagactattc 720
aaaggacaac cttctgtggt gcatgttcac atcaagtgtc actcgatgaa ggacttgcct 780
gaaccagaag acgaaattgc acagtggtgc agagatcagt ttgtggctaa ggatgcactg 840
ttagacaaac acatagctgc agacactttc cccggtcaga aagaacagaa cattggccgt 900
cccataaagt ctcttgcagt ggttgtatca tgggcatgcc tactaactct tggagcaatg 960
aagttcttac actggtcaaa cctcttttct tcgtggaaag gcatcgcatt atcagccttt 1020
ggtctaggca tcatcactct ctgtatgcag atcctgatcc gctcctctca gtcggagcgt 1080
tcaacacctg ccaaagtcgc tccagccaag ccaaaggaca atcaccagtc aggaccatcc 1140
tcccaaacag aagtggagga gaagcagaag taa 1173
<210> 58
<211> 390
<212> PRT
<213> 欧洲油菜
<400> 58
Met Ala Met Ala Ala Ala Val Ile Val Pro Leu Gly Ile Leu Phe Phe
1 5 10 15
Ile Ser Gly Leu Val Val Asn Leu Leu Gln Ala Val Cys Tyr Val Leu
20 25 30
Val Arg Pro Met Ser Lys Asn Thr Tyr Arg Lys Ile Asn Arg Val Val
35 40 45
Ala Glu Thr Leu Trp Leu Glu Leu Val Trp Ile Val Asp Trp Trp Ala
50 55 60
Gly Val Lys Ile Gln Val Phe Ala Asp Asp Glu Thr Phe Asn Arg Met
65 70 75 80
Gly Lys Glu His Ala Leu Val Val Cys Asn His Arg Ser Asp Ile Asp
85 90 95
Trp Leu Val Gly Trp Ile Leu Ala Gln Arg Ser Gly Cys Leu Gly Ser
100 105 110
Ala Leu Ala Val Met Lys Lys Ser Ser Lys Phe Leu Pro Val Ile Gly
115 120 125
Trp Ser Met Trp Phe Ser Glu Tyr Leu Phe Leu Glu Arg Asn Trp Ala
130 135 140
Lys Asp Glu Ser Thr Leu Gln Ser Gly Leu Gln Arg Leu Asn Asp Phe
145 150 155 160
Pro Arg Pro Phe Trp Leu Ala Leu Phe Val Glu Gly Thr Arg Phe Thr
165 170 175
Glu Ala Lys Leu Lys Ala Ala Gln Glu Tyr Ala Ala Ser Ser Glu Leu
180 185 190
Pro Val Pro Arg Asn Val Leu Ile Pro Arg Thr Lys Gly Phe Val Ser
195 200 205
Ala Val Ser Asn Met Arg Ser Phe Val Pro Ala Ile Tyr Asp Met Thr
210 215 220
Val Ala Ile Pro Lys Thr Ser Pro Pro Pro Thr Met Leu Arg Leu Phe
225 230 235 240
Lys Gly Gln Pro Ser Val Val His Val His Ile Lys Cys His Ser Met
245 250 255
Lys Asp Leu Pro Glu Pro Glu Asp Glu Ile Ala Gln Trp Cys Arg Asp
260 265 270
Gln Phe Val Ala Lys Asp Ala Leu Leu Asp Lys His Ile Ala Ala Asp
275 280 285
Thr Phe Pro Gly Gln Lys Glu Gln Asn Ile Gly Arg Pro Ile Lys Ser
290 295 300
Leu Ala Val Val Val Ser Trp Ala Cys Leu Leu Thr Leu Gly Ala Met
305 310 315 320
Lys Phe Leu His Trp Ser Asn Leu Phe Ser Ser Trp Lys Gly Ile Ala
325 330 335
Leu Ser Ala Phe Gly Leu Gly Ile Ile Thr Leu Cys Met Gln Ile Leu
340 345 350
Ile Arg Ser Ser Gln Ser Glu Arg Ser Thr Pro Ala Lys Val Ala Pro
355 360 365
Ala Lys Pro Lys Asp Asn His Gln Ser Gly Pro Ser Ser Gln Thr Glu
370 375 380
Val Glu Glu Lys Gln Lys
385 390
<210> 59
<211> 1152
<212> DNA
<213> 大豆
<400> 59
atgactgcag ttgtggtcgt cccattgggc cttctcttct tcgcttccgg cctcatcgtt 60
aacctcattc aggcaatatg ctacgtggtt gtgcggccgg tgtcgaagaa tctgtaccgg 120
cggatgaaca gggtggtggc ggagctgctc tggctggagc tcgtgtggat tattgattgg 180
tgggccggtg ttaaggttca agtattcaca gatcctgaaa cctttcgttc aatgggtaaa 240
gagcatgctc ttgttatctc caatcacaga agtgacattg attggcttgt tggatgggtt 300
ttagctcagc gttcaggttg ccttggcagc actcttgctg tgatgaagaa atcatcgaag 360
tttctaccgg tcattggttg gtcaatgtgg ttttctgaat atcttttcct ggagagaagt 420
tgggccaagg atgaacgcac attaaagtca ggcctacagc aactgaggga tttccctctt 480
cccttttggt tggctctctt tgtagaagga actcgcttta cacaggccaa actattagct 540
gctcaggagt atgcagcctc agctggattg cctgttccaa gaaatgtttt gattccaaga 600
actaagggtt ttgtttcagc agtaaaccat atgcgctctt ttgttcctgc catttatgat 660
gtaactgtgg caattcccaa gagttcacct gctcctacaa tgctaagact cttcagaggg 720
aaatcttcag tggttcatgt gcatattaag cggcatgcaa tgaaggattt gccagaagaa 780
gatgaagctg ttgctcaatg gtgtcgagat atgtttgtgg ctaaggatac attgttagac 840
aaacatatag ctgaggacac attcagtgat caagagctgc aggataccgg tcgacccata 900
aagtctctag tggttgtcat atcgtgggca tgtgtcgttg ttatgggggt cgtaaagttc 960
ctccaatggt catcactact atcctcctgg aagggtgttg cattttcagc atttggtttg 1020
ggagttgtta ctctactcat gcacatcttg atcatgttct cacaatctga gcgttcaacc 1080
cctaccaagg ttgcccctgc aaagtccaag aatagcgaac aactggaggc tagggataac 1140
aaacaagact ag 1152
<210> 60
<211> 383
<212> PRT
<213> 大豆
<400> 60
Met Thr Ala Val Val Val Val Pro Leu Gly Leu Leu Phe Phe Ala Ser
1 5 10 15
Gly Leu Ile Val Asn Leu Ile Gln Ala Ile Cys Tyr Val Val Val Arg
20 25 30
Pro Val Ser Lys Asn Leu Tyr Arg Arg Met Asn Arg Val Val Ala Glu
35 40 45
Leu Leu Trp Leu Glu Leu Val Trp Ile Ile Asp Trp Trp Ala Gly Val
50 55 60
Lys Val Gln Val Phe Thr Asp Pro Glu Thr Phe Arg Ser Met Gly Lys
65 70 75 80
Glu His Ala Leu Val Ile Ser Asn His Arg Ser Asp Ile Asp Trp Leu
85 90 95
Val Gly Trp Val Leu Ala Gln Arg Ser Gly Cys Leu Gly Ser Thr Leu
100 105 110
Ala Val Met Lys Lys Ser Ser Lys Phe Leu Pro Val Ile Gly Trp Ser
115 120 125
Met Trp Phe Ser Glu Tyr Leu Phe Leu Glu Arg Ser Trp Ala Lys Asp
130 135 140
Glu Arg Thr Leu Lys Ser Gly Leu Gln Gln Leu Arg Asp Phe Pro Leu
145 150 155 160
Pro Phe Trp Leu Ala Leu Phe Val Glu Gly Thr Arg Phe Thr Gln Ala
165 170 175
Lys Leu Leu Ala Ala Gln Glu Tyr Ala Ala Ser Ala Gly Leu Pro Val
180 185 190
Pro Arg Asn Val Leu Ile Pro Arg Thr Lys Gly Phe Val Ser Ala Val
195 200 205
Asn His Met Arg Ser Phe Val Pro Ala Ile Tyr Asp Val Thr Val Ala
210 215 220
Ile Pro Lys Ser Ser Pro Ala Pro Thr Met Leu Arg Leu Phe Arg Gly
225 230 235 240
Lys Ser Ser Val Val His Val His Ile Lys Arg His Ala Met Lys Asp
245 250 255
Leu Pro Glu Glu Asp Glu Ala Val Ala Gln Trp Cys Arg Asp Met Phe
260 265 270
Val Ala Lys Asp Thr Leu Leu Asp Lys His Ile Ala Glu Asp Thr Phe
275 280 285
Ser Asp Gln Glu Leu Gln Asp Thr Gly Arg Pro Ile Lys Ser Leu Val
290 295 300
Val Val Ile Ser Trp Ala Cys Val Val Val Met Gly Val Val Lys Phe
305 310 315 320
Leu Gln Trp Ser Ser Leu Leu Ser Ser Trp Lys Gly Val Ala Phe Ser
325 330 335
Ala Phe Gly Leu Gly Val Val Thr Leu Leu Met His Ile Leu Ile Met
340 345 350
Phe Ser Gln Ser Glu Arg Ser Thr Pro Thr Lys Val Ala Pro Ala Lys
355 360 365
Ser Lys Asn Ser Glu Gln Leu Glu Ala Arg Asp Asn Lys Gln Asp
370 375 380
<210> 61
<211> 849
<212> DNA
<213> 解脂耶氏酵母
<400> 61
atgtccgttg catccaagct cgtcttctac gtccgcgccg ccatcgccgt ggtcatcttt 60
gccgcctgtg ccacctacgg cgtgctggcg tccaccattc tcaccgccat cggcaagcag 120
ggcctggccc aatggaccgt tgccagagcc ttctactact cggtgcgcat cttcctgggt 180
atcagcatca agctgcgtag ccggcaggtg accggaaccg ccggtctgga tgcctccaag 240
atccaggtcg ccaacaccac caagcccatt gacgacatca ccaaacacct gccccgacca 300
tgcattctga tttccaacca ccagaacgaa atggacattc tggtgctcgg tcgcatcttc 360
ccccagtact gctccgtcac cgccaaaaag gccctcaagt ggtaccctct gctgggccag 420
ttcatggcgc tgtccggcac catcttcctg gaccgaaagg accgaaccaa gtccgtgcag 480
accctcggcg gcgccgtcaa gaccatccag agcggcaacg gaggcaaggg ccagagcgtc 540
ttcatgttcc ccgagggaac ccgatcctac tccaaggacg tcggcatcat gcccttcaag 600
aagggctgtt tccacctggc ggtccagtcg ggcgctccca ttgtccccgt ggtggtccag 660
aacacctccc gaatgttttc tttcggccga ggcaagctgg acgccggaga gatccttgtc 720
gacgtcctga gccccattga gaccaagggt ctggacgcca gcaacgtcga cgctctcatg 780
gccaccactt ataaggccat gtgcgagact gccgaccaga ttggctacgc tggccagaag 840
actcagtag 849
<210> 62
<211> 282
<212> PRT
<213> 解脂耶氏酵母
<400> 62
Met Ser Val Ala Ser Lys Leu Val Phe Tyr Val Arg Ala Ala Ile Ala
1 5 10 15
Val Val Ile Phe Ala Ala Cys Ala Thr Tyr Gly Val Leu Ala Ser Thr
20 25 30
Ile Leu Thr Ala Ile Gly Lys Gln Gly Leu Ala Gln Trp Thr Val Ala
35 40 45
Arg Ala Phe Tyr Tyr Ser Val Arg Ile Phe Leu Gly Ile Ser Ile Lys
50 55 60
Leu Arg Ser Arg Gln Val Thr Gly Thr Ala Gly Leu Asp Ala Ser Lys
65 70 75 80
Ile Gln Val Ala Asn Thr Thr Lys Pro Ile Asp Asp Ile Thr Lys His
85 90 95
Leu Pro Arg Pro Cys Ile Leu Ile Ser Asn His Gln Asn Glu Met Asp
100 105 110
Ile Leu Val Leu Gly Arg Ile Phe Pro Gln Tyr Cys Ser Val Thr Ala
115 120 125
Lys Lys Ala Leu Lys Trp Tyr Pro Leu Leu Gly Gln Phe Met Ala Leu
130 135 140
Ser Gly Thr Ile Phe Leu Asp Arg Lys Asp Arg Thr Lys Ser Val Gln
145 150 155 160
Thr Leu Gly Gly Ala Val Lys Thr Ile Gln Ser Gly Asn Gly Gly Lys
165 170 175
Gly Gln Ser Val Phe Met Phe Pro Glu Gly Thr Arg Ser Tyr Ser Lys
180 185 190
Asp Val Gly Ile Met Pro Phe Lys Lys Gly Cys Phe His Leu Ala Val
195 200 205
Gln Ser Gly Ala Pro Ile Val Pro Val Val Val Gln Asn Thr Ser Arg
210 215 220
Met Phe Ser Phe Gly Arg Gly Lys Leu Asp Ala Gly Glu Ile Leu Val
225 230 235 240
Asp Val Leu Ser Pro Ile Glu Thr Lys Gly Leu Asp Ala Ser Asn Val
245 250 255
Asp Ala Leu Met Ala Thr Thr Tyr Lys Ala Met Cys Glu Thr Ala Asp
260 265 270
Gln Ile Gly Tyr Ala Gly Gln Lys Thr Gln
275 280
<210> 63
<211> 906
<212> DNA
<213> 拟南芥
<400> 63
atgtcagccg ccgcagctga aaccgacgtc tctctccgtc gcagatctaa ctctcttaac 60
ggaaaccaca ctaacggcgt cgccattgac ggaaccctag acaacaacaa ccgtcgcgtc 120
ggagatacaa acactcacat ggatatatct gctaagaaaa ctgacaacgg ctacgccaat 180
ggtgtcggag gaggaggatg gagaagcaaa gcgtcgttca cgacgtggac ggcgcgtgat 240
atcgtctacg tggtgagata ccattggata ccgtgcatgt tcgctgccgg acttctgttc 300
ttcatgggcg tggagtacac gcttcagatg attcccgcga gatctgagcc gttcgatctt 360
gggtttgtgg tcacgcgctc tttgaatcgc gtattagcat cttcaccgga tcttaacact 420
gttttagccg cactaaacac ggtgttcgta gggatgcaaa caacgtatat tgtatggaca 480
tggttagtgg aaggacgagc acgagccacc atcgcggctt tattcatgtt cacttgtcgc 540
ggcattctcg gctactctac tcagcttcct ctccctcagg actttctagg atcaggggtt 600
gattttccgg tgggaaatgt ctctttcttc ctcttcttct ctggccatgt cgccggctcg 660
atgatcgcat cattggacat gagaagaatg cagaggttga gacttgcaat ggtctttgac 720
atcctcaatg tattacagtc gatcagactg ctcggtacaa gaggacatta cacaatcgac 780
cttgcggttg gagttggcgc tgggattctc ttcgactcat tggccggaaa gtacgaagag 840
atgatgagca agagacattt aggcactggt tttagtttga tttcgaaaga ctctctagtc 900
aattaa 906
<210> 64
<211> 301
<212> PRT
<213> 拟南芥
<400> 64
Met Ser Ala Ala Ala Ala Glu Thr Asp Val Ser Leu Arg Arg Arg Ser
1 5 10 15
Asn Ser Leu Asn Gly Asn His Thr Asn Gly Val Ala Ile Asp Gly Thr
20 25 30
Leu Asp Asn Asn Asn Arg Arg Val Gly Asp Thr Asn Thr His Met Asp
35 40 45
Ile Ser Ala Lys Lys Thr Asp Asn Gly Tyr Ala Asn Gly Val Gly Gly
50 55 60
Gly Gly Trp Arg Ser Lys Ala Ser Phe Thr Thr Trp Thr Ala Arg Asp
65 70 75 80
Ile Val Tyr Val Val Arg Tyr His Trp Ile Pro Cys Met Phe Ala Ala
85 90 95
Gly Leu Leu Phe Phe Met Gly Val Glu Tyr Thr Leu Gln Met Ile Pro
100 105 110
Ala Arg Ser Glu Pro Phe Asp Leu Gly Phe Val Val Thr Arg Ser Leu
115 120 125
Asn Arg Val Leu Ala Ser Ser Pro Asp Leu Asn Thr Val Leu Ala Ala
130 135 140
Leu Asn Thr Val Phe Val Gly Met Gln Thr Thr Tyr Ile Val Trp Thr
145 150 155 160
Trp Leu Val Glu Gly Arg Ala Arg Ala Thr Ile Ala Ala Leu Phe Met
165 170 175
Phe Thr Cys Arg Gly Ile Leu Gly Tyr Ser Thr Gln Leu Pro Leu Pro
180 185 190
Gln Asp Phe Leu Gly Ser Gly Val Asp Phe Pro Val Gly Asn Val Ser
195 200 205
Phe Phe Leu Phe Phe Ser Gly His Val Ala Gly Ser Met Ile Ala Ser
210 215 220
Leu Asp Met Arg Arg Met Gln Arg Leu Arg Leu Ala Met Val Phe Asp
225 230 235 240
Ile Leu Asn Val Leu Gln Ser Ile Arg Leu Leu Gly Thr Arg Gly His
245 250 255
Tyr Thr Ile Asp Leu Ala Val Gly Val Gly Ala Gly Ile Leu Phe Asp
260 265 270
Ser Leu Ala Gly Lys Tyr Glu Glu Met Met Ser Lys Arg His Leu Gly
275 280 285
Thr Gly Phe Ser Leu Ile Ser Lys Asp Ser Leu Val Asn
290 295 300
<210> 65
<211> 840
<212> DNA
<213> 向日葵
<400> 65
atggggacaa tcacaatgaa cgccgataag atccaccaac gttccatacc acaaacaacc 60
accctcctca aaacatcaaa caacgtttcc attaacaatt tcaatctcaa aaagaccaac 120
aattctacga ccacatgggg atcgctagcg gacgcgtcgt ttttacaatg gacaacatcg 180
gatgtgttcg gtttgtttaa gtaccacccc gtcccgtgtt tctttgctgt ttcgttactg 240
ttttttatgg gtgtggagta tacgctgcgg atgataccgc cctcgtcacc cccgtttgat 300
attgggttcg ttgccactgc gtatttgcat cgtgtcctcg tcgctagtcc gacgctaaac 360
acagttctcg cagggcttaa tacggtgttt gtggggatgc aaacaaccta tataataggg 420
acgtgggtca ttgaagggcg gccgagagcc acagtcgcgg cgctgttcat gtttacttgc 480
agagggattt tgggttattc gacacagctt cctttgcctg agggattctt gggttcggga 540
gcagactttc cggtaggcaa tgtgtccttc ttcttgtttt attctggtca tgtagcagga 600
tcagtaattg catcgtttga tatgaagaga atgcataggt gggagttggc ctttctattc 660
gacaccttga accttttgca ggtggtgcgg ttactaagca caaggggaca ctacaccatt 720
gacttagccg tcggagtagg tgccggcatg ttgtttgatt ccctcgccgg aaaatacatg 780
aagaaggcta tggctgaagg aggtgatgtc ttgtatacaa acctacctca cttatcttaa 840
<210> 66
<211> 279
<212> PRT
<213> 向日葵
<400> 66
Met Gly Thr Ile Thr Met Asn Ala Asp Lys Ile His Gln Arg Ser Ile
1 5 10 15
Pro Gln Thr Thr Thr Leu Leu Lys Thr Ser Asn Asn Val Ser Ile Asn
20 25 30
Asn Phe Asn Leu Lys Lys Thr Asn Asn Ser Thr Thr Thr Trp Gly Ser
35 40 45
Leu Ala Asp Ala Ser Phe Leu Gln Trp Thr Thr Ser Asp Val Phe Gly
50 55 60
Leu Phe Lys Tyr His Pro Val Pro Cys Phe Phe Ala Val Ser Leu Leu
65 70 75 80
Phe Phe Met Gly Val Glu Tyr Thr Leu Arg Met Ile Pro Pro Ser Ser
85 90 95
Pro Pro Phe Asp Ile Gly Phe Val Ala Thr Ala Tyr Leu His Arg Val
100 105 110
Leu Val Ala Ser Pro Thr Leu Asn Thr Val Leu Ala Gly Leu Asn Thr
115 120 125
Val Phe Val Gly Met Gln Thr Thr Tyr Ile Ile Gly Thr Trp Val Ile
130 135 140
Glu Gly Arg Pro Arg Ala Thr Val Ala Ala Leu Phe Met Phe Thr Cys
145 150 155 160
Arg Gly Ile Leu Gly Tyr Ser Thr Gln Leu Pro Leu Pro Glu Gly Phe
165 170 175
Leu Gly Ser Gly Ala Asp Phe Pro Val Gly Asn Val Ser Phe Phe Leu
180 185 190
Phe Tyr Ser Gly His Val Ala Gly Ser Val Ile Ala Ser Phe Asp Met
195 200 205
Lys Arg Met His Arg Trp Glu Leu Ala Phe Leu Phe Asp Thr Leu Asn
210 215 220
Leu Leu Gln Val Val Arg Leu Leu Ser Thr Arg Gly His Tyr Thr Ile
225 230 235 240
Asp Leu Ala Val Gly Val Gly Ala Gly Met Leu Phe Asp Ser Leu Ala
245 250 255
Gly Lys Tyr Met Lys Lys Ala Met Ala Glu Gly Gly Asp Val Leu Tyr
260 265 270
Thr Asn Leu Pro His Leu Ser
275
<210> 67
<211> 1261
<212> DNA
<213> 亚麻荠
<400> 67
caccaaagag ttagagtgta tttaactaac aaccataaat taaaaatgtc cttctcaaaa 60
ataactattc atcactttca tatgacaatt ttctaaccca aacaagtctc ctttgaaatc 120
tccgccctcg tatctcttat ataacacatc tctcttcaaa ctcccaaaaa tgtccctctc 180
cgttaactct aacggccatg tcagtcgccg cagctaaacc cgccgtctct cgccgtcacg 240
tatctaacgg aaacaacact aacaacgtcg ccattgacga cgatcacaac caccaacgcc 300
gcatcgtcgg agataaaaac actcgaatgg agatcgctgc taagaacaac ggctacgcca 360
acggtgtcat cggaggagga ggatggagga gcaaggcgtc gttcatgacg tggacgacgc 420
gtgacgttgt ctacgtggcg agacaccatt ggataccgtg catgttcgct gccgggcttt 480
tgttcttcat gggggtcgag tacacgctcc agatgattcc cgcgagatct gagccgttcg 540
atcttgggtt cgtggccacg cgctctttga atcgcgtctt agcatcttcc ccggatctta 600
acaccgtttt agccgcacta aacacggtgt tcgtattgat gcaaacaacg tatattgtat 660
ggacatggtt agtggaagga cgagcacgag caaccatctc ggctttattc atgttcacgt 720
gtcggggcat tctcggctac tctactcagc ttcctctccc tcaggatttt ttaggatcag 780
gagttgattt tccagtggga aacgtctctt tcttcctctt cttctcgggc cacgttgccg 840
gctcgatgat cgcatcactg gacatgagga gaatgcagag gtttaagctg gcgagggttt 900
ttgacatcct caatgtatta caatcgatca ggctgctcgg tacaagagga cactacacca 960
tcgaccttgc ggttggagtt ggcgctggga ttctctttga ctcactggcc gggaagtacg 1020
aagagatgag cagaagacac cacctaggaa ctggttttag tttgatatcg aaagactctc 1080
tagtcaatta aatttgtttt ctatcaaatg tttttagttc aacacattta gttaagttga 1140
atttagtcta atgacttatt tatttcttct ttataaatgc ctaaatggtt cattgtatac 1200
cgtttgttga acctaattat cgatctaatg ttacttaaag ggttgtagcc ttgtagttgt 1260
a 1261
<210> 68
<211> 297
<212> PRT
<213> 亚麻荠
<400> 68
Met Ser Val Ala Ala Ala Lys Pro Ala Val Ser Arg Arg His Val Ser
1 5 10 15
Asn Gly Asn Asn Thr Asn Asn Val Ala Ile Asp Asp Asp His Asn His
20 25 30
Gln Arg Arg Ile Val Gly Asp Lys Asn Thr Arg Met Glu Ile Ala Ala
35 40 45
Lys Asn Asn Gly Tyr Ala Asn Gly Val Ile Gly Gly Gly Gly Trp Arg
50 55 60
Ser Lys Ala Ser Phe Met Thr Trp Thr Thr Arg Asp Val Val Tyr Val
65 70 75 80
Ala Arg His His Trp Ile Pro Cys Met Phe Ala Ala Gly Leu Leu Phe
85 90 95
Phe Met Gly Val Glu Tyr Thr Leu Gln Met Ile Pro Ala Arg Ser Glu
100 105 110
Pro Phe Asp Leu Gly Phe Val Ala Thr Arg Ser Leu Asn Arg Val Leu
115 120 125
Ala Ser Ser Pro Asp Leu Asn Thr Val Leu Ala Ala Leu Asn Thr Val
130 135 140
Phe Val Leu Met Gln Thr Thr Tyr Ile Val Trp Thr Trp Leu Val Glu
145 150 155 160
Gly Arg Ala Arg Ala Thr Ile Ser Ala Leu Phe Met Phe Thr Cys Arg
165 170 175
Gly Ile Leu Gly Tyr Ser Thr Gln Leu Pro Leu Pro Gln Asp Phe Leu
180 185 190
Gly Ser Gly Val Asp Phe Pro Val Gly Asn Val Ser Phe Phe Leu Phe
195 200 205
Phe Ser Gly His Val Ala Gly Ser Met Ile Ala Ser Leu Asp Met Arg
210 215 220
Arg Met Gln Arg Phe Lys Leu Ala Arg Val Phe Asp Ile Leu Asn Val
225 230 235 240
Leu Gln Ser Ile Arg Leu Leu Gly Thr Arg Gly His Tyr Thr Ile Asp
245 250 255
Leu Ala Val Gly Val Gly Ala Gly Ile Leu Phe Asp Ser Leu Ala Gly
260 265 270
Lys Tyr Glu Glu Met Ser Arg Arg His His Leu Gly Thr Gly Phe Ser
275 280 285
Leu Ile Ser Lys Asp Ser Leu Val Asn
290 295
<210> 69
<211> 1204
<212> DNA
<213> 欧洲油菜
<400> 69
atgaactttt gtagcccaaa caaccttcct ttccttccac aagtttcata atatctctta 60
tataacccat atctccaagc ctctcgaaat gttcttctcc gttaaatcta acggtcatgt 120
caactacaac aatcgtccct ctccgtcgca gttctaactc tctcaatgaa taccacacta 180
acgcagtcgc ctttgacgga atcgtcgggt caacaagtac tagccaaatg gaggagattg 240
ttacgcaaac cgacgactgc tacgccaatc acaacggaga tggagggaga agcaaggcat 300
cgtttatgac gtggaggatg tgcaatcctg tccaggtggc gagagtccat tggataccgt 360
gtttgctagc ggtaggagtt ctgttcttca cgggcgtaga ggagtacatg ctccagatga 420
ttccggcgag ttctgagccg ttcgatattg gttttgtggc gacgcgctct ctgtatcgac 480
tcttggcttc ttcaccggat cttaataccg ttttagctgc tctcaacacg gtgtttgtag 540
ggatgcaaac gacgtatatt ttatggacat ggttggtgga aggacgacca cgagcgacca 600
tctcggcttg cttcatgttt acttgtcgtg gcattcttgg ttactctact cagctccctc 660
ttcctcagga ttttctagga tcaggggtag attttccggt aggaaacgtc tcgttcttcc 720
tcttctactc aggccatgtc gcagggtcga cgatagcatc cttggatatg aggagaatga 780
agaggttgag acttgccttg ctttttgaca tcctcaatgt attacaatcg atcaggcttc 840
tcgggacgag aggacaatac acgatcgatc tcgctgtcgg agttggcgct ggggttctct 900
ttgactcact ggctggaaaa tacgaagaga tgatgagcaa gagacgcaat gtaggcaatg 960
gttttagttt gatttcgtct cgctagttat taatttttgt tttttttttt ttatgttttt 1020
agtctggaca tatttaattt agttgaaatc taatgactta aatttgcttt ctttcaaaat 1080
gctctaactg acggacctaa ctaaatgtgt acgttattgt gtagttacca tagaggtttc 1140
gtattgtctt gagcctgata ttttgatttt agagctcgtt tatacggtag ctaataataa 1200
aaaa 1204
<210> 70
<211> 289
<212> PRT
<213> 欧洲油菜
<400> 70
Met Ser Thr Thr Thr Ile Val Pro Leu Arg Arg Ser Ser Asn Ser Leu
1 5 10 15
Asn Glu Tyr His Thr Asn Ala Val Ala Phe Asp Gly Ile Val Gly Ser
20 25 30
Thr Ser Thr Ser Gln Met Glu Glu Ile Val Thr Gln Thr Asp Asp Cys
35 40 45
Tyr Ala Asn His Asn Gly Asp Gly Gly Arg Ser Lys Ala Ser Phe Met
50 55 60
Thr Trp Arg Met Cys Asn Pro Val Gln Val Ala Arg Val His Trp Ile
65 70 75 80
Pro Cys Leu Leu Ala Val Gly Val Leu Phe Phe Thr Gly Val Glu Glu
85 90 95
Tyr Met Leu Gln Met Ile Pro Ala Ser Ser Glu Pro Phe Asp Ile Gly
100 105 110
Phe Val Ala Thr Arg Ser Leu Tyr Arg Leu Leu Ala Ser Ser Pro Asp
115 120 125
Leu Asn Thr Val Leu Ala Ala Leu Asn Thr Val Phe Val Gly Met Gln
130 135 140
Thr Thr Tyr Ile Leu Trp Thr Trp Leu Val Glu Gly Arg Pro Arg Ala
145 150 155 160
Thr Ile Ser Ala Cys Phe Met Phe Thr Cys Arg Gly Ile Leu Gly Tyr
165 170 175
Ser Thr Gln Leu Pro Leu Pro Gln Asp Phe Leu Gly Ser Gly Val Asp
180 185 190
Phe Pro Val Gly Asn Val Ser Phe Phe Leu Phe Tyr Ser Gly His Val
195 200 205
Ala Gly Ser Thr Ile Ala Ser Leu Asp Met Arg Arg Met Lys Arg Leu
210 215 220
Arg Leu Ala Leu Leu Phe Asp Ile Leu Asn Val Leu Gln Ser Ile Arg
225 230 235 240
Leu Leu Gly Thr Arg Gly Gln Tyr Thr Ile Asp Leu Ala Val Gly Val
245 250 255
Gly Ala Gly Val Leu Phe Asp Ser Leu Ala Gly Lys Tyr Glu Glu Met
260 265 270
Met Ser Lys Arg Arg Asn Val Gly Asn Gly Phe Ser Leu Ile Ser Ser
275 280 285
Arg
<210> 71
<211> 804
<212> DNA
<213> 大豆
<400> 71
atgaatggcg gcgctgaggc ctccctcaat cacaggcgca aacaccaaac agctcccgcc 60
gacggcgcta aaggcgttaa ggtagcaaac ggagccatgg ggaagccgtc ctcttccaag 120
cactcctgcg gcgcgtcgtt catgaaatgg accgtggctg acgctgtcca cgtggtgacg 180
caccattgga tgccgtgctt gttcgcattg gggcttctct tcttcatggc cgtggagtac 240
acgcttctca tggtgccgcc gtcgtcgccg cccttcgacc ttggcttcat cgccacacgc 300
tccctccacg cgctcctcga gtcgtcgccg aatctcaaca cgctcttcgc cgggctcaat 360
acggtgtttg tggggatgca aacgagttat atcttatgga cgtggctgat tgaaggacgc 420
cccagagcca caatttcagc attgttcatg ttcacatgcc gtgggatttt aggctactcc 480
acccagctcc cattgcctca ggggtttttg ggctcgggtg tggacttccc tgttgggaac 540
gtgtcttttt tcttgttttt ttctgggcac gttgcagggt cagtgattgc ttcattggac 600
atgaggagga tgcagaggtg ggaactggct tggacttttg atgtgctcaa tgttttgcaa 660
gctgtgaggt tgctgggtac aagaggccat tacactattg atttggccgt aggggttggt 720
gctggaattc tctttgattc tttagctggc aagtacgaag atagcaaaag gaatggtgct 780
ctcaaacaca atttgattgc gtga 804
<210> 72
<211> 267
<212> PRT
<213> 大豆
<400> 72
Met Asn Gly Gly Ala Glu Ala Ser Leu Asn His Arg Arg Lys His Gln
1 5 10 15
Thr Ala Pro Ala Asp Gly Ala Lys Gly Val Lys Val Ala Asn Gly Ala
20 25 30
Met Gly Lys Pro Ser Ser Ser Lys His Ser Cys Gly Ala Ser Phe Met
35 40 45
Lys Trp Thr Val Ala Asp Ala Val His Val Val Thr His His Trp Met
50 55 60
Pro Cys Leu Phe Ala Leu Gly Leu Leu Phe Phe Met Ala Val Glu Tyr
65 70 75 80
Thr Leu Leu Met Val Pro Pro Ser Ser Pro Pro Phe Asp Leu Gly Phe
85 90 95
Ile Ala Thr Arg Ser Leu His Ala Leu Leu Glu Ser Ser Pro Asn Leu
100 105 110
Asn Thr Leu Phe Ala Gly Leu Asn Thr Val Phe Val Gly Met Gln Thr
115 120 125
Ser Tyr Ile Leu Trp Thr Trp Leu Ile Glu Gly Arg Pro Arg Ala Thr
130 135 140
Ile Ser Ala Leu Phe Met Phe Thr Cys Arg Gly Ile Leu Gly Tyr Ser
145 150 155 160
Thr Gln Leu Pro Leu Pro Gln Gly Phe Leu Gly Ser Gly Val Asp Phe
165 170 175
Pro Val Gly Asn Val Ser Phe Phe Leu Phe Phe Ser Gly His Val Ala
180 185 190
Gly Ser Val Ile Ala Ser Leu Asp Met Arg Arg Met Gln Arg Trp Glu
195 200 205
Leu Ala Trp Thr Phe Asp Val Leu Asn Val Leu Gln Ala Val Arg Leu
210 215 220
Leu Gly Thr Arg Gly His Tyr Thr Ile Asp Leu Ala Val Gly Val Gly
225 230 235 240
Ala Gly Ile Leu Phe Asp Ser Leu Ala Gly Lys Tyr Glu Asp Ser Lys
245 250 255
Arg Asn Gly Ala Leu Lys His Asn Leu Ile Ala
260 265
<210> 73
<211> 412
<212> PRT
<213> 拟南芥
<400> 73
Met Val Ala Thr Ser Ala Thr Ser Ser Phe Phe Pro Val Pro Ser Ser
1 5 10 15
Ser Leu Asp Pro Asn Gly Lys Gly Asn Lys Ile Gly Ser Thr Asn Leu
20 25 30
Ala Gly Leu Asn Ser Ala Pro Asn Ser Gly Arg Met Lys Val Lys Pro
35 40 45
Asn Ala Gln Ala Pro Pro Lys Ile Asn Gly Lys Lys Val Gly Leu Pro
50 55 60
Gly Ser Val Asp Ile Val Arg Thr Asp Thr Glu Thr Ser Ser His Pro
65 70 75 80
Ala Pro Arg Thr Phe Ile Asn Gln Leu Pro Asp Trp Ser Met Leu Leu
85 90 95
Ala Ala Ile Thr Thr Ile Phe Leu Ala Ala Glu Lys Gln Trp Met Met
100 105 110
Leu Asp Trp Lys Pro Arg Arg Ser Asp Met Leu Val Asp Pro Phe Gly
115 120 125
Ile Gly Arg Ile Val Gln Asp Gly Leu Val Phe Arg Gln Asn Phe Ser
130 135 140
Ile Arg Ser Tyr Glu Ile Gly Ala Asp Arg Ser Ala Ser Ile Glu Thr
145 150 155 160
Val Met Asn His Leu Gln Glu Thr Ala Leu Asn His Val Lys Thr Ala
165 170 175
Gly Leu Leu Gly Asp Gly Phe Gly Ser Thr Pro Glu Met Phe Lys Lys
180 185 190
Asn Leu Ile Trp Val Val Thr Arg Met Gln Val Val Val Asp Lys Tyr
195 200 205
Pro Thr Trp Gly Asp Val Val Glu Val Asp Thr Trp Val Ser Gln Ser
210 215 220
Gly Lys Asn Gly Met Arg Arg Asp Trp Leu Val Arg Asp Cys Asn Thr
225 230 235 240
Gly Glu Thr Leu Thr Arg Ala Ser Ser Val Trp Val Met Met Asn Lys
245 250 255
Leu Thr Arg Arg Leu Ser Lys Ile Pro Glu Glu Val Arg Gly Glu Ile
260 265 270
Glu Pro Tyr Phe Val Asn Ser Asp Pro Val Leu Ala Glu Asp Ser Arg
275 280 285
Lys Leu Thr Lys Ile Asp Asp Lys Thr Ala Asp Tyr Val Arg Ser Gly
290 295 300
Leu Thr Pro Arg Trp Ser Asp Leu Asp Val Asn Gln His Val Asn Asn
305 310 315 320
Val Lys Tyr Ile Gly Trp Ile Leu Glu Ser Ala Pro Val Gly Ile Met
325 330 335
Glu Arg Gln Lys Leu Lys Ser Met Thr Leu Glu Tyr Arg Arg Glu Cys
340 345 350
Gly Arg Asp Ser Val Leu Gln Ser Leu Thr Ala Val Thr Gly Cys Asp
355 360 365
Ile Gly Asn Leu Ala Thr Ala Gly Asp Val Glu Cys Gln His Leu Leu
370 375 380
Arg Leu Gln Asp Gly Ala Glu Val Val Arg Gly Arg Thr Glu Trp Ser
385 390 395 400
Ser Lys Thr Pro Thr Thr Thr Trp Gly Thr Ala Pro
405 410
<210> 74
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> AGPAT1 引物 1
<400> 74
caccatggat ttatggcctg gtgc 24
<210> 75
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> AGPAT1 引物 2
<400> 75
tcatcctcct ccacctgg 18
<210> 76
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> AGPAT1 引物 3
<400> 76
cggaattcat ggatttatgg cctggtgc 28
<210> 77
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> AGPAT1 引物 4
<400> 77
gctctagatc atcctcctcc acctgg 26
<210> 78
<211> 1626
<212> DNA
<213> 拟南芥
<400> 78
atggtgggtg cgtcttcctc ttacgcatct ccgttatgta cctggtttgt tgctgcttgc 60
atgtccgtct ctcacggtgg aggagatagc cgtcaggctg ttgctcttca atctggtggg 120
cggagtcggc gaaggaggca gcttagcaaa tgctctgtcg cttctggatc cgctagcatt 180
caggctctcg tcacttcttg tttggatttt ggtccttgta ctcactacaa caacaacaat 240
gcattgtctt ctctctttgg atcgaatagt gtttctttga atcgaaacca gaggagattg 300
aatcgtgctg ctagctccgg tggagccatg gcagtgatgg agatggaaaa ggaagctgcg 360
gttaacaaga aaccacctac ggagcagcgt cgagttgtag tgacaggcat gggagttgaa 420
acatcattgg gtcatgaccc acataccttc tatgagaatt tgctacaagg caacagtggt 480
attagccaga ttgaaaattt tgattgttct gaatttccta cgcgaattgc gggagagatc 540
aaaagcttct cgactgaagg atgggttgct ccaaaacttt ctaaaaggat ggacaaattc 600
atgctctatc ttctcacagc tggtaagaaa gctttggctg atggtggggt tactgatgaa 660
gtaatggcag agtttgacaa aaccaaatgt ggagttttga ttggctcggc aatgggagga 720
atgaaggtct tttacgatgc tattgaagct ctgagaatct cttacaagaa gatgaatcct 780
ttttgtgtac cttttgcgac aacaaacatg ggttctgcta tgcttgccat ggatctggga 840
tggatggggc caaactattc tatttcaact gcttgtgcca caagcaactt ttgcattctg 900
aattcagcaa accacattat taaaggtgaa gctgatgtaa tgctctgtgg tggctcagat 960
gcagttatta ttccaatagg gttgggaggt tttgttgcat gccgggctct ttcacaaagg 1020
aataatgatc ccacaaaagc ttcacgtcct tgggatacca atcgagatgg tttcgtgatg 1080
ggagagggag ctggagttct acttttggaa gaactcgagc atgctaagaa aagaggtgca 1140
actatctacg cagagttcct cggtgggagt ttcacatgtg atgcctatca catgaccgag 1200
cctcaccctg atggggctgg tgttattctc tgtattgaga gagcgttagc tagtgctggg 1260
atttccaagg aacaaataaa ttacataaat gcacatgcaa cctcaacgca tgctggagat 1320
attaaggaat accaagccct tgctcactgt tttggccaaa atcctgagct taaggtaaat 1380
tccacaaaat ctatgattgg acacttgctg ggagctgctg gggccgtgga ggctgttgca 1440
actgtgcagg cgatacggac cggatgggtt catccaaata tcaacctcga gaatccagac 1500
agtggagtgg atacaaagct gctggtgggt cctaagaagg agagactgga cattaaagca 1560
gccttgtcaa attcattcgg gtttggtggt cataactcca gcatcatttt tgctccttac 1620
aagtga 1626
<210> 79
<211> 541
<212> PRT
<213> 拟南芥
<400> 79
Met Val Gly Ala Ser Ser Ser Tyr Ala Ser Pro Leu Cys Thr Trp Phe
1 5 10 15
Val Ala Ala Cys Met Ser Val Ser His Gly Gly Gly Asp Ser Arg Gln
20 25 30
Ala Val Ala Leu Gln Ser Gly Gly Arg Ser Arg Arg Arg Arg Gln Leu
35 40 45
Ser Lys Cys Ser Val Ala Ser Gly Ser Ala Ser Ile Gln Ala Leu Val
50 55 60
Thr Ser Cys Leu Asp Phe Gly Pro Cys Thr His Tyr Asn Asn Asn Asn
65 70 75 80
Ala Leu Ser Ser Leu Phe Gly Ser Asn Ser Val Ser Leu Asn Arg Asn
85 90 95
Gln Arg Arg Leu Asn Arg Ala Ala Ser Ser Gly Gly Ala Met Ala Val
100 105 110
Met Glu Met Glu Lys Glu Ala Ala Val Asn Lys Lys Pro Pro Thr Glu
115 120 125
Gln Arg Arg Val Val Val Thr Gly Met Gly Val Glu Thr Ser Leu Gly
130 135 140
His Asp Pro His Thr Phe Tyr Glu Asn Leu Leu Gln Gly Asn Ser Gly
145 150 155 160
Ile Ser Gln Ile Glu Asn Phe Asp Cys Ser Glu Phe Pro Thr Arg Ile
165 170 175
Ala Gly Glu Ile Lys Ser Phe Ser Thr Glu Gly Trp Val Ala Pro Lys
180 185 190
Leu Ser Lys Arg Met Asp Lys Phe Met Leu Tyr Leu Leu Thr Ala Gly
195 200 205
Lys Lys Ala Leu Ala Asp Gly Gly Val Thr Asp Glu Val Met Ala Glu
210 215 220
Phe Asp Lys Thr Lys Cys Gly Val Leu Ile Gly Ser Ala Met Gly Gly
225 230 235 240
Met Lys Val Phe Tyr Asp Ala Ile Glu Ala Leu Arg Ile Ser Tyr Lys
245 250 255
Lys Met Asn Pro Phe Cys Val Pro Phe Ala Thr Thr Asn Met Gly Ser
260 265 270
Ala Met Leu Ala Met Asp Leu Gly Trp Met Gly Pro Asn Tyr Ser Ile
275 280 285
Ser Thr Ala Cys Ala Thr Ser Asn Phe Cys Ile Leu Asn Ser Ala Asn
290 295 300
His Ile Ile Lys Gly Glu Ala Asp Val Met Leu Cys Gly Gly Ser Asp
305 310 315 320
Ala Val Ile Ile Pro Ile Gly Leu Gly Gly Phe Val Ala Cys Arg Ala
325 330 335
Leu Ser Gln Arg Asn Asn Asp Pro Thr Lys Ala Ser Arg Pro Trp Asp
340 345 350
Thr Asn Arg Asp Gly Phe Val Met Gly Glu Gly Ala Gly Val Leu Leu
355 360 365
Leu Glu Glu Leu Glu His Ala Lys Lys Arg Gly Ala Thr Ile Tyr Ala
370 375 380
Glu Phe Leu Gly Gly Ser Phe Thr Cys Asp Ala Tyr His Met Thr Glu
385 390 395 400
Pro His Pro Asp Gly Ala Gly Val Ile Leu Cys Ile Glu Arg Ala Leu
405 410 415
Ala Ser Ala Gly Ile Ser Lys Glu Gln Ile Asn Tyr Ile Asn Ala His
420 425 430
Ala Thr Ser Thr His Ala Gly Asp Ile Lys Glu Tyr Gln Ala Leu Ala
435 440 445
His Cys Phe Gly Gln Asn Pro Glu Leu Lys Val Asn Ser Thr Lys Ser
450 455 460
Met Ile Gly His Leu Leu Gly Ala Ala Gly Ala Val Glu Ala Val Ala
465 470 475 480
Thr Val Gln Ala Ile Arg Thr Gly Trp Val His Pro Asn Ile Asn Leu
485 490 495
Glu Asn Pro Asp Ser Gly Val Asp Thr Lys Leu Leu Val Gly Pro Lys
500 505 510
Lys Glu Arg Leu Asp Ile Lys Ala Ala Leu Ser Asn Ser Phe Gly Phe
515 520 525
Gly Gly His Asn Ser Ser Ile Ile Phe Ala Pro Tyr Lys
530 535 540
<210> 80
<211> 1152
<212> DNA
<213> 拟南芥
<400> 80
atgggtgcag gtggaagaat gccggttcct acttcttcca agaaatcgga aaccgacacc 60
acaaagcgtg tgccgtgcga gaaaccgcct ttctcggtgg gagatctgaa gaaagcaatc 120
ccgccgcatt gtttcaaacg ctcaatccct cgctctttct cctaccttat cagtgacatc 180
attatagcct catgcttcta ctacgtcgcc accaattact tctctctcct ccctcagcct 240
ctctcttact tggcttggcc actctattgg gcctgtcaag gctgtgtcct aactggtatc 300
tgggtcatag cccacgaatg cggtcaccac gcattcagcg actaccaatg gctggatgac 360
acagttggtc ttatcttcca ttccttcctc ctcgtccctt acttctcctg gaagtatagt 420
catcgccgtc accattccaa cactggatcc ctcgaaagag atgaagtatt tgtcccaaag 480
cagaaatcag caatcaagtg gtacgggaaa tacctcaaca accctcttgg acgcatcatg 540
atgttaaccg tccagtttgt cctcgggtgg cccttgtact tagcctttaa cgtctctggc 600
agaccgtatg acgggttcgc ttgccatttc ttccccaacg ctcccatcta caatgaccga 660
gaacgcctcc agatatacct ctctgatgcg ggtattctag ccgtctgttt tggtctttac 720
cgttacgctg ctgcacaagg gatggcctcg atgatctgcc tctacggagt accgcttctg 780
atagtgaatg cgttcctcgt cttgatcact tacttgcagc acactcatcc ctcgttgcct 840
cactacgatt catcagagtg ggactggctc aggggagctt tggctaccgt agacagagac 900
tacggaatct tgaacaaggt gttccacaac attacagaca cacacgtggc tcatcacctg 960
ttctcgacaa tgccgcatta taacgcaatg gaagctacaa aggcgataaa gccaattctg 1020
ggagactatt accagttcga tggaacaccg tggtatgtag cgatgtatag ggaggcaaag 1080
gagtgtatct atgtagaacc ggacagggaa ggtgacaaga aaggtgtgta ctggtacaac 1140
aataagttat ga 1152
<210> 81
<211> 383
<212> PRT
<213> 拟南芥
<400> 81
Met Gly Ala Gly Gly Arg Met Pro Val Pro Thr Ser Ser Lys Lys Ser
1 5 10 15
Glu Thr Asp Thr Thr Lys Arg Val Pro Cys Glu Lys Pro Pro Phe Ser
20 25 30
Val Gly Asp Leu Lys Lys Ala Ile Pro Pro His Cys Phe Lys Arg Ser
35 40 45
Ile Pro Arg Ser Phe Ser Tyr Leu Ile Ser Asp Ile Ile Ile Ala Ser
50 55 60
Cys Phe Tyr Tyr Val Ala Thr Asn Tyr Phe Ser Leu Leu Pro Gln Pro
65 70 75 80
Leu Ser Tyr Leu Ala Trp Pro Leu Tyr Trp Ala Cys Gln Gly Cys Val
85 90 95
Leu Thr Gly Ile Trp Val Ile Ala His Glu Cys Gly His His Ala Phe
100 105 110
Ser Asp Tyr Gln Trp Leu Asp Asp Thr Val Gly Leu Ile Phe His Ser
115 120 125
Phe Leu Leu Val Pro Tyr Phe Ser Trp Lys Tyr Ser His Arg Arg His
130 135 140
His Ser Asn Thr Gly Ser Leu Glu Arg Asp Glu Val Phe Val Pro Lys
145 150 155 160
Gln Lys Ser Ala Ile Lys Trp Tyr Gly Lys Tyr Leu Asn Asn Pro Leu
165 170 175
Gly Arg Ile Met Met Leu Thr Val Gln Phe Val Leu Gly Trp Pro Leu
180 185 190
Tyr Leu Ala Phe Asn Val Ser Gly Arg Pro Tyr Asp Gly Phe Ala Cys
195 200 205
His Phe Phe Pro Asn Ala Pro Ile Tyr Asn Asp Arg Glu Arg Leu Gln
210 215 220
Ile Tyr Leu Ser Asp Ala Gly Ile Leu Ala Val Cys Phe Gly Leu Tyr
225 230 235 240
Arg Tyr Ala Ala Ala Gln Gly Met Ala Ser Met Ile Cys Leu Tyr Gly
245 250 255
Val Pro Leu Leu Ile Val Asn Ala Phe Leu Val Leu Ile Thr Tyr Leu
260 265 270
Gln His Thr His Pro Ser Leu Pro His Tyr Asp Ser Ser Glu Trp Asp
275 280 285
Trp Leu Arg Gly Ala Leu Ala Thr Val Asp Arg Asp Tyr Gly Ile Leu
290 295 300
Asn Lys Val Phe His Asn Ile Thr Asp Thr His Val Ala His His Leu
305 310 315 320
Phe Ser Thr Met Pro His Tyr Asn Ala Met Glu Ala Thr Lys Ala Ile
325 330 335
Lys Pro Ile Leu Gly Asp Tyr Tyr Gln Phe Asp Gly Thr Pro Trp Tyr
340 345 350
Val Ala Met Tyr Arg Glu Ala Lys Glu Cys Ile Tyr Val Glu Pro Asp
355 360 365
Arg Glu Gly Asp Lys Lys Gly Val Tyr Trp Tyr Asn Asn Lys Leu
370 375 380
<210> 82
<211> 1521
<212> DNA
<213> 拟南芥
<400> 82
atgacgtccg ttaacgttaa gctcctttac cgttacgtct taaccaactt tttcaacctc 60
tgtttgttcc cgttaacggc gttcctcgcc ggaaaagcct ctcggcttac cataaacgat 120
ctccacaact tcctttccta tctccaacac aaccttataa cagtaacttt actctttgct 180
ttcactgttt tcggtttggt tctctacatc gtaacccgac ccaatccggt ttatctcgtt 240
gactactcgt gttaccttcc accaccgcat ctcaaagtta gtgtctctaa agtcatggat 300
attttctacc aaataagaaa agctgatact tcttcacgga acgtggcatg tgatgatccg 360
tcctcgctcg atttcctgag gaagattcaa gagcgttcag gtctaggtga tgagacgtac 420
agtcctgagg gactcattca cgtaccaccg cggaagactt ttgcagcgtc acgtgaagag 480
acagagaagg ttatcatcgg tgcgctcgaa aatctattcg agaacaccaa agttaaccct 540
agagagattg gtatacttgt ggtgaactca agcatgttta atccaactcc ttcgctatcc 600
gctatggtcg ttaatacttt caagctccga agcaacatca aaagctttaa tctaggagga 660
atgggttgta gtgctggtgt tattgccatt gatttggcta aagacttgtt gcatgttcat 720
aaaaacactt atgctcttgt ggtgagcact gagaacatca cacaaggcat ttatgctgga 780
gaaaatagat caatgatggt tagcaattgc ttgtttcgtg ttggtggggc cgcgattttg 840
ctctctaaca agtcgggaga ccggagacgg tccaagtaca agctagttca cacggtccga 900
acgcatactg gagctgatga caagtctttt cgatgtgtgc aacaagaaga cgatgagagc 960
ggcaaaatcg gagtttgtct gtcaaaggac ataaccaatg ttgcggggac aacacttacg 1020
aaaaatatag caacattggg tccgttgatt cttcctttaa gcgaaaagtt tctttttttc 1080
gctaccttcg tcgccaagaa acttctaaag gataaaatca agcattacta tgttccggat 1140
ttcaagcttg ctgttgacca tttctgtatt catgccggag gcagagccgt gatcgatgag 1200
ctagagaaga acttaggact atcgccgatc gatgtggagg catctagatc aacgttacat 1260
agatttggga atacttcatc tagctcaatt tggtatgaat tagcatacat agaggcaaag 1320
ggaagaatga agaaagggaa taaagcttgg cagattgctt taggatcagg gtttaagtgt 1380
aatagtgcgg tttgggtggc tctacgcaat gtcaaggcat cggcaaatag tccttggcaa 1440
cattgcatcg atagatatcc ggttaaaatt gattctgatt tgtcaaagtc aaagactcat 1500
gtccaaaacg gtcggtccta a 1521
<210> 83
<211> 506
<212> PRT
<213> 拟南芥
<400> 83
Met Thr Ser Val Asn Val Lys Leu Leu Tyr Arg Tyr Val Leu Thr Asn
1 5 10 15
Phe Phe Asn Leu Cys Leu Phe Pro Leu Thr Ala Phe Leu Ala Gly Lys
20 25 30
Ala Ser Arg Leu Thr Ile Asn Asp Leu His Asn Phe Leu Ser Tyr Leu
35 40 45
Gln His Asn Leu Ile Thr Val Thr Leu Leu Phe Ala Phe Thr Val Phe
50 55 60
Gly Leu Val Leu Tyr Ile Val Thr Arg Pro Asn Pro Val Tyr Leu Val
65 70 75 80
Asp Tyr Ser Cys Tyr Leu Pro Pro Pro His Leu Lys Val Ser Val Ser
85 90 95
Lys Val Met Asp Ile Phe Tyr Gln Ile Arg Lys Ala Asp Thr Ser Ser
100 105 110
Arg Asn Val Ala Cys Asp Asp Pro Ser Ser Leu Asp Phe Leu Arg Lys
115 120 125
Ile Gln Glu Arg Ser Gly Leu Gly Asp Glu Thr Tyr Ser Pro Glu Gly
130 135 140
Leu Ile His Val Pro Pro Arg Lys Thr Phe Ala Ala Ser Arg Glu Glu
145 150 155 160
Thr Glu Lys Val Ile Ile Gly Ala Leu Glu Asn Leu Phe Glu Asn Thr
165 170 175
Lys Val Asn Pro Arg Glu Ile Gly Ile Leu Val Val Asn Ser Ser Met
180 185 190
Phe Asn Pro Thr Pro Ser Leu Ser Ala Met Val Val Asn Thr Phe Lys
195 200 205
Leu Arg Ser Asn Ile Lys Ser Phe Asn Leu Gly Gly Met Gly Cys Ser
210 215 220
Ala Gly Val Ile Ala Ile Asp Leu Ala Lys Asp Leu Leu His Val His
225 230 235 240
Lys Asn Thr Tyr Ala Leu Val Val Ser Thr Glu Asn Ile Thr Gln Gly
245 250 255
Ile Tyr Ala Gly Glu Asn Arg Ser Met Met Val Ser Asn Cys Leu Phe
260 265 270
Arg Val Gly Gly Ala Ala Ile Leu Leu Ser Asn Lys Ser Gly Asp Arg
275 280 285
Arg Arg Ser Lys Tyr Lys Leu Val His Thr Val Arg Thr His Thr Gly
290 295 300
Ala Asp Asp Lys Ser Phe Arg Cys Val Gln Gln Glu Asp Asp Glu Ser
305 310 315 320
Gly Lys Ile Gly Val Cys Leu Ser Lys Asp Ile Thr Asn Val Ala Gly
325 330 335
Thr Thr Leu Thr Lys Asn Ile Ala Thr Leu Gly Pro Leu Ile Leu Pro
340 345 350
Leu Ser Glu Lys Phe Leu Phe Phe Ala Thr Phe Val Ala Lys Lys Leu
355 360 365
Leu Lys Asp Lys Ile Lys His Tyr Tyr Val Pro Asp Phe Lys Leu Ala
370 375 380
Val Asp His Phe Cys Ile His Ala Gly Gly Arg Ala Val Ile Asp Glu
385 390 395 400
Leu Glu Lys Asn Leu Gly Leu Ser Pro Ile Asp Val Glu Ala Ser Arg
405 410 415
Ser Thr Leu His Arg Phe Gly Asn Thr Ser Ser Ser Ser Ile Trp Tyr
420 425 430
Glu Leu Ala Tyr Ile Glu Ala Lys Gly Arg Met Lys Lys Gly Asn Lys
435 440 445
Ala Trp Gln Ile Ala Leu Gly Ser Gly Phe Lys Cys Asn Ser Ala Val
450 455 460
Trp Val Ala Leu Arg Asn Val Lys Ala Ser Ala Asn Ser Pro Trp Gln
465 470 475 480
His Cys Ile Asp Arg Tyr Pro Val Lys Ile Asp Ser Asp Leu Ser Lys
485 490 495
Ser Lys Thr His Val Gln Asn Gly Arg Ser
500 505
<210> 84
<211> 1260
<212> DNA
<213> 解脂耶氏酵母
<400> 84
atggattcga ccacgcagac caacaccggc accggcaagg tggccgtgca gccccccacg 60
gccttcatta agcccattga gaaggtgtcc gagcccgtct acgacacctt tggcaacgag 120
ttcactcctc cagactactc tatcaaggat attctggatg ccattcccca ggagtgctac 180
aagcggtcct acgttaagtc ctactcgtac gtggcccgag actgcttctt tatcgccgtt 240
tttgcctaca tggcctacgc gtacctgcct cttattccct cggcttccgg ccgagctgtg 300
gcctgggcca tgtactccat tgtccagggt ctgtttggca ccggtctgtg ggttcttgcc 360
cacgagtgtg gccactctgc tttctccgac tctaacaccg tcaacaacgt caccggatgg 420
gttctgcact cctccatgct ggtcccttac tacgcctgga agctgaccca ctccatgcac 480
cacaagtcca ctggtcacct cacccgtgat atggtgtttg tgcccaagga ccgaaaggag 540
tttatggaga accgaggcgc ccatgactgg tctgagcttg ctgaggacgc tcccctcatg 600
accctctacg gcctcatcac ccagcaggtg tttggatggc ctctgtatct gctgtctaac 660
gttaccggac agaagtaccc caagctcaac aaatgggctg tcaaccactt caaccccaac 720
gccccgctgt ttgagaagaa ggactggttc aacatctgga tctctaacgt cggtattggt 780
atcaccatgt ccgtcatcgc atactccatc aaccgatggg gcctggcttc cgtcaccctc 840
tactacctga tcccctacct gtgggtcaac cactggctcg tggccatcac ctacctgcag 900
cacaccgacc ccactctgcc ccactaccac gccgaccagt ggaacttcac ccgaggagcc 960
gccgccacca tcgaccgaga gtttggcttc atcggctcct tctgcttcca tgacatcatc 1020
gagacccacg ttctgcacca ctacgtgtct cgaattccct tctacaacgc ccgaatcgcc 1080
actgagaaga tcaagaaggt catgggcaag cactaccgac acgacgacac caacttcatc 1140
aagtctcttt acactgtcgc ccgaacctgc cagtttgttg aaggtaagga aggcattcag 1200
atgtttagaa acgtcaatgg agtcggagtt gctcctgacg gcctgccttc taaaaagtag 1260
<210> 85
<211> 419
<212> PRT
<213> 解脂耶氏酵母
<400> 85
Met Asp Ser Thr Thr Gln Thr Asn Thr Gly Thr Gly Lys Val Ala Val
1 5 10 15
Gln Pro Pro Thr Ala Phe Ile Lys Pro Ile Glu Lys Val Ser Glu Pro
20 25 30
Val Tyr Asp Thr Phe Gly Asn Glu Phe Thr Pro Pro Asp Tyr Ser Ile
35 40 45
Lys Asp Ile Leu Asp Ala Ile Pro Gln Glu Cys Tyr Lys Arg Ser Tyr
50 55 60
Val Lys Ser Tyr Ser Tyr Val Ala Arg Asp Cys Phe Phe Ile Ala Val
65 70 75 80
Phe Ala Tyr Met Ala Tyr Ala Tyr Leu Pro Leu Ile Pro Ser Ala Ser
85 90 95
Gly Arg Ala Val Ala Trp Ala Met Tyr Ser Ile Val Gln Gly Leu Phe
100 105 110
Gly Thr Gly Leu Trp Val Leu Ala His Glu Cys Gly His Ser Ala Phe
115 120 125
Ser Asp Ser Asn Thr Val Asn Asn Val Thr Gly Trp Val Leu His Ser
130 135 140
Ser Met Leu Val Pro Tyr Tyr Ala Trp Lys Leu Thr His Ser Met His
145 150 155 160
His Lys Ser Thr Gly His Leu Thr Arg Asp Met Val Phe Val Pro Lys
165 170 175
Asp Arg Lys Glu Phe Met Glu Asn Arg Gly Ala His Asp Trp Ser Glu
180 185 190
Leu Ala Glu Asp Ala Pro Leu Met Thr Leu Tyr Gly Leu Ile Thr Gln
195 200 205
Gln Val Phe Gly Trp Pro Leu Tyr Leu Leu Ser Asn Val Thr Gly Gln
210 215 220
Lys Tyr Pro Lys Leu Asn Lys Trp Ala Val Asn His Phe Asn Pro Asn
225 230 235 240
Ala Pro Leu Phe Glu Lys Lys Asp Trp Phe Asn Ile Trp Ile Ser Asn
245 250 255
Val Gly Ile Gly Ile Thr Met Ser Val Ile Ala Tyr Ser Ile Asn Arg
260 265 270
Trp Gly Leu Ala Ser Val Thr Leu Tyr Tyr Leu Ile Pro Tyr Leu Trp
275 280 285
Val Asn His Trp Leu Val Ala Ile Thr Tyr Leu Gln His Thr Asp Pro
290 295 300
Thr Leu Pro His Tyr His Ala Asp Gln Trp Asn Phe Thr Arg Gly Ala
305 310 315 320
Ala Ala Thr Ile Asp Arg Glu Phe Gly Phe Ile Gly Ser Phe Cys Phe
325 330 335
His Asp Ile Ile Glu Thr His Val Leu His His Tyr Val Ser Arg Ile
340 345 350
Pro Phe Tyr Asn Ala Arg Ile Ala Thr Glu Lys Ile Lys Lys Val Met
355 360 365
Gly Lys His Tyr Arg His Asp Asp Thr Asn Phe Ile Lys Ser Leu Tyr
370 375 380
Thr Val Ala Arg Thr Cys Gln Phe Val Glu Gly Lys Glu Gly Ile Gln
385 390 395 400
Met Phe Arg Asn Val Asn Gly Val Gly Val Ala Pro Asp Gly Leu Pro
405 410 415
Ser Lys Lys
<210> 86
<211> 978
<212> DNA
<213> 解脂耶氏酵母
<400> 86
atgagcgccg tccctattga attcaacgtc ccctccgtgg accgaccctt tggtatctac 60
ctctgggcca tctttgacca ggcctgggag aagcttttcg gctggcccgc gtcctctttc 120
attttcgtgc gaaatgaccc caacatcccc ttttcctcta cccctcccgt gatcattgcc 180
atcattgtgt actacattgt catctttggc ggccgagagg tgatgcgaaa cctgtctccc 240
atccgactca actggctctt ccagatccac aacatcttcc tcacccttct gtccggtatg 300
ctcctcctcc tcctcgttga gcagctcttc cccatcattg tccgacaggg tatcctctac 360
gccatctgcg actacggatc ttggactcag cccattgtct tctgctacta cctcaactac 420
ctgaccaagt actttgagct gatcgacacc gttttccttg tgctgcgaaa gaagaagctg 480
actttcctcc acacctacca ccatggtgcc actgctcttc tgtgctacac ccagctcatt 540
ggtaagacct cggtctcttg ggtccccatc acccttaacc tgtttgtcca cgttgtcatg 600
tacttctact acttcctggc tgcgcgaggt atccgagtgt ggtggaagga gtgggtcacc 660
cggctccaga tcatccagtt cgttatcgat cttggatttg tctactttgc ctcttacacc 720
tacttcacct ctacctactg gccctggatg cccaacatgg gctcttgtgc cggcgaggag 780
tttgctgcta tttacggctg tggtctgctg acctcttacc tcttcctctt catcgccttc 840
tacatcaact cttaccgaaa gccctcttcc aagggacctt ccaagcctgt tgttgctgtc 900
gatggccctg ttggcggcgt caacgcccag actggtgctt ctcgaggcca gaccactacc 960
cgatctcgac gagcataa 978
<210> 87
<211> 325
<212> PRT
<213> 解脂耶氏酵母
<400> 87
Met Ser Ala Val Pro Ile Glu Phe Asn Val Pro Ser Val Asp Arg Pro
1 5 10 15
Phe Gly Ile Tyr Leu Trp Ala Ile Phe Asp Gln Ala Trp Glu Lys Leu
20 25 30
Phe Gly Trp Pro Ala Ser Ser Phe Ile Phe Val Arg Asn Asp Pro Asn
35 40 45
Ile Pro Phe Ser Ser Thr Pro Pro Val Ile Ile Ala Ile Ile Val Tyr
50 55 60
Tyr Ile Val Ile Phe Gly Gly Arg Glu Val Met Arg Asn Leu Ser Pro
65 70 75 80
Ile Arg Leu Asn Trp Leu Phe Gln Ile His Asn Ile Phe Leu Thr Leu
85 90 95
Leu Ser Gly Met Leu Leu Leu Leu Leu Val Glu Gln Leu Phe Pro Ile
100 105 110
Ile Val Arg Gln Gly Ile Leu Tyr Ala Ile Cys Asp Tyr Gly Ser Trp
115 120 125
Thr Gln Pro Ile Val Phe Cys Tyr Tyr Leu Asn Tyr Leu Thr Lys Tyr
130 135 140
Phe Glu Leu Ile Asp Thr Val Phe Leu Val Leu Arg Lys Lys Lys Leu
145 150 155 160
Thr Phe Leu His Thr Tyr His His Gly Ala Thr Ala Leu Leu Cys Tyr
165 170 175
Thr Gln Leu Ile Gly Lys Thr Ser Val Ser Trp Val Pro Ile Thr Leu
180 185 190
Asn Leu Phe Val His Val Val Met Tyr Phe Tyr Tyr Phe Leu Ala Ala
195 200 205
Arg Gly Ile Arg Val Trp Trp Lys Glu Trp Val Thr Arg Leu Gln Ile
210 215 220
Ile Gln Phe Val Ile Asp Leu Gly Phe Val Tyr Phe Ala Ser Tyr Thr
225 230 235 240
Tyr Phe Thr Ser Thr Tyr Trp Pro Trp Met Pro Asn Met Gly Ser Cys
245 250 255
Ala Gly Glu Glu Phe Ala Ala Ile Tyr Gly Cys Gly Leu Leu Thr Ser
260 265 270
Tyr Leu Phe Leu Phe Ile Ala Phe Tyr Ile Asn Ser Tyr Arg Lys Pro
275 280 285
Ser Ser Lys Gly Pro Ser Lys Pro Val Val Ala Val Asp Gly Pro Val
290 295 300
Gly Gly Val Asn Ala Gln Thr Gly Ala Ser Arg Gly Gln Thr Thr Thr
305 310 315 320
Arg Ser Arg Arg Ala
325
<210> 88
<211> 1185
<212> DNA
<213> 解脂耶氏酵母
<400> 88
atgggcgtat tcattaaaca ggagcagctt ccggctctca agaagtacaa gtactccgcc 60
gaggatcact cgttcatctc caacaacatt ctgcgcccct tctggcgaca gtttgtcaaa 120
atcttccctc tgtggatggc ccccaacatg gtgactctgt tgggcttctt ctttgtcatt 180
gtgaacttca tcaccatgct cattgttgat cccacccacg accgcgagcc tcccagatgg 240
gtctacctca cctacgctct gggtctgttc ctttaccaga catttgatgc ctgtgacgga 300
tcccatgccc gacgaactgg ccagagtgga ccccttggag agctgtttga ccactgtgtc 360
gacgccatga atacctctct gattctcacg gtggtggtgt ccaccaccca tatgggatat 420
aacatgaagc tgctgattgt gcagattgcc gctctcggaa acttctacct gtcgacctgg 480
gagacctacc ataccggaac tctgtacctt tctggcttct ctggtcctgt tgaaggtatc 540
ttgattctgg tggctctttt cgtcctcacc ttcttcactg gtcccaacgt gtacgctctg 600
accgtctacg aggctcttcc cgaatccatc acttcgctgc tgcctgccag cttcctggac 660
gtcaccatca cccagatcta cattggattc ggagtgctgg gcatggtgtt caacatctac 720
ggcgcctgcg gaaacgtgat caagtactac aacaacaagg gcaagagcgc tctccccgcc 780
attctcggaa tcgccccctt tggcatcttc tacgtcggcg tctttgcctg ggcccatgtt 840
gctcctctgc ttctctccaa gtacgccatc gtctatctgt ttgccattgg ggctgccttt 900
gccatgcaag tcggccagat gattcttgcc catctcgtgc ttgctccctt cccccactgg 960
aacgtgctgc tcttcttccc ctttgtggga ctggcagtgc actacattgc acccgtgttt 1020
ggctgggacg ccgatatcgt gtcggttaac actctcttca cctgttttgg cgccaccctc 1080
tccatttacg ccttctttgt gcttgagatc atcgacgaga tcaccaacta cctcgatatc 1140
tggtgtctgc gaatcaagta ccctcaggag aagaagactg agtaa 1185
<210> 89
<211> 394
<212> PRT
<213> 解脂耶氏酵母
<400> 89
Met Gly Val Phe Ile Lys Gln Glu Gln Leu Pro Ala Leu Lys Lys Tyr
1 5 10 15
Lys Tyr Ser Ala Glu Asp His Ser Phe Ile Ser Asn Asn Ile Leu Arg
20 25 30
Pro Phe Trp Arg Gln Phe Val Lys Ile Phe Pro Leu Trp Met Ala Pro
35 40 45
Asn Met Val Thr Leu Leu Gly Phe Phe Phe Val Ile Val Asn Phe Ile
50 55 60
Thr Met Leu Ile Val Asp Pro Thr His Asp Arg Glu Pro Pro Arg Trp
65 70 75 80
Val Tyr Leu Thr Tyr Ala Leu Gly Leu Phe Leu Tyr Gln Thr Phe Asp
85 90 95
Ala Cys Asp Gly Ser His Ala Arg Arg Thr Gly Gln Ser Gly Pro Leu
100 105 110
Gly Glu Leu Phe Asp His Cys Val Asp Ala Met Asn Thr Ser Leu Ile
115 120 125
Leu Thr Val Val Val Ser Thr Thr His Met Gly Tyr Asn Met Lys Leu
130 135 140
Leu Ile Val Gln Ile Ala Ala Leu Gly Asn Phe Tyr Leu Ser Thr Trp
145 150 155 160
Glu Thr Tyr His Thr Gly Thr Leu Tyr Leu Ser Gly Phe Ser Gly Pro
165 170 175
Val Glu Gly Ile Leu Ile Leu Val Ala Leu Phe Val Leu Thr Phe Phe
180 185 190
Thr Gly Pro Asn Val Tyr Ala Leu Thr Val Tyr Glu Ala Leu Pro Glu
195 200 205
Ser Ile Thr Ser Leu Leu Pro Ala Ser Phe Leu Asp Val Thr Ile Thr
210 215 220
Gln Ile Tyr Ile Gly Phe Gly Val Leu Gly Met Val Phe Asn Ile Tyr
225 230 235 240
Gly Ala Cys Gly Asn Val Ile Lys Tyr Tyr Asn Asn Lys Gly Lys Ser
245 250 255
Ala Leu Pro Ala Ile Leu Gly Ile Ala Pro Phe Gly Ile Phe Tyr Val
260 265 270
Gly Val Phe Ala Trp Ala His Val Ala Pro Leu Leu Leu Ser Lys Tyr
275 280 285
Ala Ile Val Tyr Leu Phe Ala Ile Gly Ala Ala Phe Ala Met Gln Val
290 295 300
Gly Gln Met Ile Leu Ala His Leu Val Leu Ala Pro Phe Pro His Trp
305 310 315 320
Asn Val Leu Leu Phe Phe Pro Phe Val Gly Leu Ala Val His Tyr Ile
325 330 335
Ala Pro Val Phe Gly Trp Asp Ala Asp Ile Val Ser Val Asn Thr Leu
340 345 350
Phe Thr Cys Phe Gly Ala Thr Leu Ser Ile Tyr Ala Phe Phe Val Leu
355 360 365
Glu Ile Ile Asp Glu Ile Thr Asn Tyr Leu Asp Ile Trp Cys Leu Arg
370 375 380
Ile Lys Tyr Pro Gln Glu Lys Lys Thr Glu
385 390
<210> 90
<211> 1332
<212> DNA
<213> 解脂耶氏酵母
<400> 90
atgggtaaaa gcccctctat gattgggact tgcggttggg ttgggctggg tacaaacaca 60
tgctatgctc atcagctcag ccgtgccatg ggccatcaac caacttttct cccgacccta 120
atcccctcca cactactaat gccaggattc caccacatca accccgacag tctagtgcat 180
ctaaagtcgt accagtacag aagcgtcgat aaatcctacc tgtctaagta catcctgaac 240
ccatggtgga cttacgcagc aaccttcatg cctgattggc tggctcccaa cgccatcact 300
ctcatcggtg tctccggcat gctcctgtca atcttcttca ccgtctggta cacccccgag 360
ctcaccggcg acggtccctc ctggatctac ttcttctctg ccttttcgct cttcttctac 420
cagaccatgg acaatattga cggcaagcag gcccgtcgaa ctggctcttc ctctcctctg 480
ggagagctgt ttgaccacgg aatcgactcc ctcaactgca cctacggcgg aattgtcaac 540
tgcggtgctg tggcccttgg ttccacctcc tacggaggcc tcatggttct gtccacctgt 600
attggtatgt acttttctac ctgggagact tactacaccc acactcttta cctgggagtt 660
gttaacggcc ccaccgaagg gctagtggtg gccctgtcta ccatgctaat ctccggcttc 720
atgggcaccg acatctggaa ggaggatgcc gttgaggtgc tccccttcct ctccttcatg 780
gttcccgagt acctcaagct caacgagttc tgggtctggg tcgtcatgtt caccctcgtg 840
gtgctgcacg tgcctttctg cgtgtggaac gtctattggg cctgcaagga ggacgatgtg 900
cctttctccg aggctctggt gggcctcctg ccctttggcg tggccggagg agctgcctac 960
gtgtggctgc agagtcctta ttccactgtg ctggttgaca accaccttgt gctatttgga 1020
ctgaccgcct cgtgggtctt tggccggctg accaccggtg tgattctcaa ccacctcacc 1080
aagctcgagt tccctctgtg gaactccacc ctgattcctc ttctgggagc caccgttctg 1140
ttctacctgc tccctgccct gggtctactg ccccaggaca atccccactt cgagactctg 1200
tatctgtggg gcttctttgt ctacgctgcc gcgaactttt tgacctgggc tgttaacacc 1260
atcaatgtca tctgttccta ccttggcatc cgatgtctgt ctctgcgacc cgtggacaac 1320
aagaccaact ag 1332
<210> 91
<211> 443
<212> PRT
<213> 解脂耶氏酵母
<400> 91
Met Gly Lys Ser Pro Ser Met Ile Gly Thr Cys Gly Trp Val Gly Leu
1 5 10 15
Gly Thr Asn Thr Cys Tyr Ala His Gln Leu Ser Arg Ala Met Gly His
20 25 30
Gln Pro Thr Phe Leu Pro Thr Leu Ile Pro Ser Thr Leu Leu Met Pro
35 40 45
Gly Phe His His Ile Asn Pro Asp Ser Leu Val His Leu Lys Ser Tyr
50 55 60
Gln Tyr Arg Ser Val Asp Lys Ser Tyr Leu Ser Lys Tyr Ile Leu Asn
65 70 75 80
Pro Trp Trp Thr Tyr Ala Ala Thr Phe Met Pro Asp Trp Leu Ala Pro
85 90 95
Asn Ala Ile Thr Leu Ile Gly Val Ser Gly Met Leu Leu Ser Ile Phe
100 105 110
Phe Thr Val Trp Tyr Thr Pro Glu Leu Thr Gly Asp Gly Pro Ser Trp
115 120 125
Ile Tyr Phe Phe Ser Ala Phe Ser Leu Phe Phe Tyr Gln Thr Met Asp
130 135 140
Asn Ile Asp Gly Lys Gln Ala Arg Arg Thr Gly Ser Ser Ser Pro Leu
145 150 155 160
Gly Glu Leu Phe Asp His Gly Ile Asp Ser Leu Asn Cys Thr Tyr Gly
165 170 175
Gly Ile Val Asn Cys Gly Ala Val Ala Leu Gly Ser Thr Ser Tyr Gly
180 185 190
Gly Leu Met Val Leu Ser Thr Cys Ile Gly Met Tyr Phe Ser Thr Trp
195 200 205
Glu Thr Tyr Tyr Thr His Thr Leu Tyr Leu Gly Val Val Asn Gly Pro
210 215 220
Thr Glu Gly Leu Val Val Ala Leu Ser Thr Met Leu Ile Ser Gly Phe
225 230 235 240
Met Gly Thr Asp Ile Trp Lys Glu Asp Ala Val Glu Val Leu Pro Phe
245 250 255
Leu Ser Phe Met Val Pro Glu Tyr Leu Lys Leu Asn Glu Phe Trp Val
260 265 270
Trp Val Val Met Phe Thr Leu Val Val Leu His Val Pro Phe Cys Val
275 280 285
Trp Asn Val Tyr Trp Ala Cys Lys Glu Asp Asp Val Pro Phe Ser Glu
290 295 300
Ala Leu Val Gly Leu Leu Pro Phe Gly Val Ala Gly Gly Ala Ala Tyr
305 310 315 320
Val Trp Leu Gln Ser Pro Tyr Ser Thr Val Leu Val Asp Asn His Leu
325 330 335
Val Leu Phe Gly Leu Thr Ala Ser Trp Val Phe Gly Arg Leu Thr Thr
340 345 350
Gly Val Ile Leu Asn His Leu Thr Lys Leu Glu Phe Pro Leu Trp Asn
355 360 365
Ser Thr Leu Ile Pro Leu Leu Gly Ala Thr Val Leu Phe Tyr Leu Leu
370 375 380
Pro Ala Leu Gly Leu Leu Pro Gln Asp Asn Pro His Phe Glu Thr Leu
385 390 395 400
Tyr Leu Trp Gly Phe Phe Val Tyr Ala Ala Ala Asn Phe Leu Thr Trp
405 410 415
Ala Val Asn Thr Ile Asn Val Ile Cys Ser Tyr Leu Gly Ile Arg Cys
420 425 430
Leu Ser Leu Arg Pro Val Asp Asn Lys Thr Asn
435 440
Claims (36)
1.一种从非人类生物体产生甘油三酯的方法，其中所述非人类生物体经遗传修饰表达对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16：0LPAT，其中，一旦表达，C16：0LPAT定位于内质网中；所述C16：0LPAT选自：缺乏功能性叶绿体靶向序列的欧洲油菜叶绿体LPAT1；
所述非人类生物体选自拟南芥、亚麻荠和欧洲油菜。
2.根据权利要求1所述的方法，其中天然内质网溶血磷脂酸酰基转移酶ER LPAT的活性在经遗传修饰的生物体中受到抑制或阻止。
3.根据权利要求1所述的方法，其中在经遗传修饰的生物体中，二酰基甘油转化为磷脂酰胆碱并且从磷脂酰胆碱转化的活性受到抑制或阻止。
4.根据权利要求1所述的方法，其中在经遗传修饰的生物体中，磷脂酰胆碱：二酰基甘油胆碱磷酸转移酶PDCT的活性受到抑制或阻止。
5.根据权利要求1所述的方法，其中经遗传修饰的生物体表达拟南芥FATB硫酯酶。
6.根据权利要求1所述的方法，其中生物体经处理或遗传修饰以表达拟南芥FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因FAD2。
7.根据权利要求1所述的方法，其中所述生物体经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因FAE1和脂肪酸去饱和酶2基因FAD2。
8.根据权利要求1所述的方法，其中(i)所述生物体经处理或遗传修饰以表达拟南芥FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因FAD2，(ii)经遗传修饰的生物体内二酰基甘油向磷脂酰胆碱的转化受到抑制或阻止，及(iii)经遗传修饰的生物体内天然内质网溶血磷脂酸酰基转移酶ER LPAT的活性受到抑制或阻止。
9.根据权利要求1所述的方法，其中(i)所述生物体经处理或遗传修饰以表达FATB硫酯酶，并且经处理或遗传修饰以破坏脂肪酸去饱和酶2基因FAD2，(ii)经遗传修饰的生物体内二酰基甘油向磷脂酰胆碱的转化受到抑制或阻止，及(iii)经遗传修饰的生物体内天然内质网溶血磷脂酸酰基转移酶ER LPAT的活性受到抑制或阻止，及(iv)C16：0LPAT是ΔCTS-LPAT1，所述ΔCTS-LPAT1是从LPAT1中删除了101个含有N-端叶绿体靶向信号的氨基酸残基。
10.根据权利要求1所述的方法，通过抑制或阻止胆碱磷酸转移酶CPT1和/或乙醇胺磷酸转移酶EPT1的活性来抑制或阻止经遗传修饰的生物体内二酰基甘油向磷脂酰胆碱的转化。
11.根据权利要求1所述的方法，其中(i)生物体经处理或遗传修饰以破坏KASII基因FAB1、脂肪酸延长酶基因FAE1和脂肪酸去饱和酶2基因FAD2，(ii)经遗传修饰的生物体内二酰基甘油向磷脂酰胆碱的转化受到抑制或阻止，及(iii)经遗传修饰的生物体内天然内质网溶血磷脂酸酰基转移酶ER LPAT的活性受到抑制或阻止。
12.根据权利要求1-11中任一项所述的方法，其中所述甘油三酯在sn-2位包含30％以上的C16：0。
13.一种从非人类生物体产生甘油三酯的方法，其中所述非人类生物体经遗传修饰表达对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16：0LPAT；所述C16：0LPAT选自：AGPAT1人类1-酰基甘油-3-磷酸-O-酰基转移酶亚型1；所述AGPAT1是由SEQ ID NO：40的氨基酸序列组成的蛋白质；或
AGPAT1是由SEQ ID NO：26、39的核苷酸序列编码的蛋白质；所述非人类生物体选自拟南芥和欧洲油菜。
14.一种用于产生甘油三酯的非人类生物体，其中所述非人类生物体在限氮培养基中添加葡萄糖或甘油或棕榈油或葡萄糖和棕榈油的混合物或甘油和棕榈油的混合物作为碳源的培养基中培养以过表达选自CreLPAT的对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16:0LPAT，其中所述非人类生物体是解脂耶氏酵母；其中C16：0LPAT选自：
CreLPAT，所述CreLPAT是由SEQ ID NO:36的核苷酸序列编码的蛋白质。
15.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含30％以上的C16：0。
16.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含35％以上的C16：0。
17.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含40％以上的C16：0。
18.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含45％以上的C16：0。
19.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含50％以上的C16：0。
20.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含55％以上的C16：0。
21.根据权利要求14所述的非人类生物体，其中所述甘油三酯在sn-2位包含60％以上的C16：0。
22.一种用于产生甘油三酯的非人类生物体，其中所述非人类生物体在限氮培养基中添加甘油作为碳源的培养基中培养以过表达选自AGPAT1的对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16:0LPAT，其中所述非人类生物体是解脂耶氏酵母；其中C16：0LPAT选自：
AGPAT1，所述AGPAT1为人类1-酰基甘油-3-磷酸-O-酰基转移酶亚型1，所述AGPAT1是由SEQ ID NO：37的核苷酸序列编码的蛋白质。
23.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含30％以上的C16：0。
24.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含35％以上的C16：0。
25.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含40％以上的C16：0。
26.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含45％以上的C16：0。
27.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含50％以上的C16：0。
28.根据权利要求22所述的非人类生物体，其中所述甘油三酯在sn-2位包含55％以上的C16：0。
29.根据权利要求22所述的非人类生物体的细胞，其中所述非人类生物体是解脂耶氏酵母。
30.根据权利要求29所述的细胞，其中所述细胞是重组细胞。
31.经遗传修饰以表达AGPAT1的酿酒酵母，其中AGPAT1是由SEQ ID NO：40的氨基酸序列组成的蛋白质；或
AGPAT1是由SEQ ID NO：37的核苷酸序列编码的蛋白质。
32.一种从权利要求14所述的非人类生物体中提取甘油三酯的方法，包括将所述非人类生物体在限氮培养基中添加葡萄糖或甘油或棕榈油或葡萄糖和棕榈油的混合物或甘油和棕榈油的混合物作为碳源的培养基中培养过表达选自CreLPAT的对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16:0LPAT，其中所述非人类生物体是解脂耶氏酵母；其中C16:0LPAT选自：CreLPAT，所述CreLPAT是由SEQ IDNO:36的核苷酸序列编码的蛋白质。
33.一种从权利要求22所述的非人类生物体中提取甘油三酯的方法，包括将所述非人类生物体在限氮培养基中添加甘油作为碳源的培养基中培养以过表达选自AGPAT1的对C16：0-辅酶A具有特异性的溶血磷脂酸酰基转移酶C16:0LPAT，其中所述非人类生物体是解脂耶氏酵母；其中C16：0LPAT选自：
AGPAT1，所述AGPAT1为人类1-酰基甘油-3-磷酸-O-酰基转移酶亚型1，所述AGPAT1是由SEQ ID NO：37的核苷酸序列编码的蛋白质。
34.一种生产婴儿配方奶粉的方法，包括根据权利要求1至13中任一项所述的方法获得甘油三酯，并使用所述甘油三酯来生产婴儿配方奶粉。
35.一种生产婴儿配方奶粉的方法，包括根据权利要求32至33中任一项所述的方法获得甘油三酯，并使用所述甘油三酯来生产婴儿配方奶粉。
36.根据权利要求34或35所述的方法，其中所述婴儿配方奶粉包含一种或多种其它成分，其中所述一种或多种其它成分包括水、乳糖、乳化剂、益生元、益生菌、维生素和/或矿物质中的一种或多种。
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| PCT/GB2020/051875 WO2021023991A1 (en) | 2019-08-07 | 2020-08-06 | Non-human organism for producing triacylglycerol |
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