KR20220096752A - 푸코오스 전이효소를 발현하는 재조합 미생물 및 이를 이용한 2’-푸코실락토오스 제조방법 - Google Patents
푸코오스 전이효소를 발현하는 재조합 미생물 및 이를 이용한 2’-푸코실락토오스 제조방법 Download PDFInfo
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Abstract
본 발명은 2’-푸코오스 전이 효소가 도입된 미생물 및 이를 이용한 2’-푸코실락토오스의 제조방법에 관한 것으로, 본 발명의 미생물 유래 푸코실트랜스퍼아제 (fucosyltransferase)는 종래의 헬리코박터 파이로리(Helicobacter pylori) 유래 효소에 비해 안전하고 고활성을 가진다.
Description
본 발명은 푸코오스 전이효소가 발현되도록 형질전환된 재조합 미생물 및 이를 이용한 2’-푸코실락토오스 제조방법에 관한 것이다.
모유 중 함량이 가장 높은 올리고당 3가지를 포함하여 137종이 푸코실화 되어 그 비율이 약 77%이고, 남은 올리고당은 대부분 시알화 된 것(39개)으로 약 28%에 해당한다. 이 중, 특히 2’-푸코실락토오스와 3’-푸코실락토오스는 프리바이오틱 (prebiotic) 효과, 병원균 장내 부착 억제 효과 그리고 면역조절시스템 조절 효과 등을 가진다. Prebiotic 효과는 지금까지 가장 많은 연구가 진행되었는데, 모유의 모유올리고당은 유아의 장에서 초기에 우점종으로 자라는 Bifidobacterium bifidum의 생육을 선택적으로 증진시키는 반면 유해균에 의해서는 이용되지 못한다. 또한, 모유올리고당은 병원균의 장내 부착을 억제하는 것으로 알려져 있는데, 이는 장재 lectin과 결합하는 병원균의 세포벽 다당체 구조가 종종 모유올리고당의 일부 구조와 유사하기 때문이다. 따라서, 모유올리고당이 수용성 리간드 유사체를 제공하여 모유 수유 유아들이 병원균에 의한 감염에 내성을 갖는 것으로 보인다. 하지만, 여성의 약 20%는 2’-푸코실락토오스을 합성하는 푸코오스 전이효소의 변이로 인해 체내에서 이들을 제대로 합성하지 못하는 것으로 알려졌다. 이로 인하여 2’-푸코실락토오스의 산업적 생산이 필요한 실정이다.
모유올리고당의 일부는 대부분 포유동물 (영장류, 소, 돼지, 염소, 양 코끼리)의 젖에서도 소량 발견된다. 이 중 염소의 젖이 모유올리고당 조성과 가장 유사한데, 이로 인해 염소 젖 치즈 생산공정의 유청 부산물로부터 대규모 막 이용 분리 기술이 제시되었다. 또한, 최근 개발된 고체상 화학 합성법 (solid-phase synthesis)을 이용하면 하루 안에 Lewis X-Lewis Y 당을 생산할 수 있고, crystalline intermediate technology를 이용하면 2’-푸코실락토오스의 빠른 합성이 가능하게 되었다. 하지만, 위의 분리법 이나 화학적 합성법의 개발에도 불구하고 모유올리고당의 산업화를 위한 대량생산에는 몇 가지 장애물이 남아있는데, 이는 본 기술들의 낮은 입체 선택성, 낮은 총 수득율과 독성 시약의 사용 등이 식품 또는 약품 산업에 적합하지 않기 때문이다. 따라서 보다 친환경적이고 수득율이 높은 생산법이 필요하게 되며 최근 미생물에 의한 고농도의 2’-FL 생산 방법이 보고 되는 추세이다. 그러나 2’-푸코실락토오스 생합성 유전자는 Bio Safety Level 2 균주 Helicobacter pylori에서 확보된 2’-푸코실락토오스를 사용하여 주로 모유 수유 대체용 분유 및 유아용 식품에 적용 시 소비자의 인식에 부정적인 요소로 작용할 수 있으며, 국내 생산 판매 허가에 적합하지 못하다. 따라서 식품 첨가물로서 보다 안정성이 높은 유전자를 적용하여 Bio Safety Level 1 균주의 유전자 도입에 의한 2’-푸코실락토오스 생산이 필요하다.
본 발명은 식품, 화장품 및 의약품 소재인 푸코실락토오스를 생산하는 전이 효소를 가지고 있는 미생물로서, Biosafety level 1 균주, 예를 들면 바실러스 메가테리움 (bacillus megaterium) 또는 아커만시아 무시니필라 (Akkermansia muciniphila)에서 유래한 2’-퓨코오스 전이 효소를 삽입한 재조합 미생물을 제공하고, 이를 이용하여 효과적인 2'-푸코실락토오스를 생산하는 방법을 제공하기 위한 것이다.
본 발명의 일 예는 α-1,2-푸코오스 전이효소 (α-1,2-fucosyltransferase)가 발현되도록 형질전환된, 재조합 미생물에 관한 것이다. 상기 α-1,2-푸코오스 전이효소는 서열번호 1 내지 5로 이루어지는 군에서 선택된 1종 이상을 포함하는 것일 수 있다.
본 발명에서 2-푸코오스 전이 효소 활성을 가진 폴리펩티드는 서열번호 1 내지 5의 아미노산 서열에 제한되지 않고, 서열번호 1 내지 5의 아미노산 서열 중의 일부 아미노산 잔기의 치환, 삽입 또는 결핍에 의해 형성한 아미노산 서열을 포함하며, 이러한 아미노산이 수식된 단백질 또는 폴리펩티드가 2-푸코오스 전이 효소 활성을 가지고 있으면 락토오스와 GDP-fucose를 기질로 하여 푸코실락토스로 전환할 수 있다.
상기 α-1,2-푸코오스 전이효소를 암호화하는 핵산 서열은, 예를 들면 서열번호 14 내지 18로 이루어지는 군에서 선택된 1종 이상의 핵산 서열을 포함하는 것일 수 있다.
상기 재조합 미생물은 GRAS (Generally Recognized As Safe) 미생물인 것일 수 있다. 현재까지 2'-푸코실락토오스 생산에 사용되는 유전자는 Helicobacter pylori에서 확보된 2’-푸코실락토오스를 사용하므로 안정성이 좋지 않으며, 이러한 문제점을 해결하기 위해 본 발명에 따른 재조합 미생물은 Bio Safety Level 1 미생물일 수 있다. 헬리코박테 파일로리(Helicobacter pylori)에서 유래한 푸코오스 전이효소(fucosyltransferase)는 바이오세이프레벨 2 (biosafety Level 2)이기 때문에 식품 소재의 생산을 위한 효소로 적합하지 않으나, 본 발명에 따른 푸코오스 전이효소는 바이오세이프레벨 1 (biosafety level 1)이기 때문에 안정성 측면에 대한 소비자의 우려가 전혀 없는 장점이 있다.
본 발명에 따른 재조합 미생물은 helicobactor pyori 유래의 α-1,2-푸코오스 전이효소를 포함하는 대조군 대비 2’-푸코실락토오스 생산성이 현저히 상승되어, 안정성 향상 및 생산성 향상이 가능하다.
구체적으로, 본 발명의 일 예에 따른 재조합 미생물은 helicobactor pyori 유래의 α-1,2-푸코오스 전이효소를 포함하는 대조군 대비 2배 이상, 2.1배 이상, 2.5배 이상, 3배 이상, 3.5배 이상, 4배 이상, 4.5배 이상, 5배 이상, 5.5배 이상, 6배 이상, 6.5배 이상, 7배 이상, 또는 7.5배 이상의 2’-푸코실락토오스 생산성을 가지는 것일 수 있다. 따라서, 상기 재조합 미생물은 α-1,2-푸코오스 제조에 유용하게 사용될 수 있으며, α-1,2-푸코오스 생산 수율을 향상시킬 수 있다.
본 발명의 일 예에 따른 재조합 미생물은, 재조합 되기 전 2'-푸코실락토오스 생산능을 가지지 않으며, 재조합에 의해 2'-푸코실락토오스 생산능을 가지는 것일 수 있다.
상기 재조합 미생물을 형질전환하는 방법은 당업계에 공지된 모든 형질전환 방법 특별한 제한 없이 선택하여 사용할 수 있으며, 예컨대, 세균 원형질체의 융합, 전기 천공법, 추진체 포격 (projectile bombardment), 및 바이러스 벡터를 사용한 감염 등으로부터 선택된 것일 수 있다.
본 발명에 따른 재조합 미생물은 바실러스 속 (Bacillus sp.) 미생물, 코리네박테리움 속 (Corynebacterium sp.) 미생물, 대장균 속 (Escherichia sp.) 미생물, 또는 효모 (yeast)인 것일 수 있다.
구체적으로, 상기 바실러스 속 (Bacillus sp.) 미생물은 바실러스 메가테리움 (Bacillus megaterium), 바실러스 서브틸리스 (Bacillus subtilis), 바실러스 세레우스 (Bacillus cereus), 바실러스 코아귤런스 (Bacillus coagulans), 바실러스 리체니포르미스 (Bacillus licheniformis), 또는 바실러스 스테아로테르모필루스 (Bacillus stearothermophilus)인 것일 수 있다.
구체적으로, 상기 코리네박테리움 속 (Corynebacterium sp.) 미생물은 코리네박테리움 글루타미쿰 (Corynebacterium glutatamicum)인 것일 수 있다.
구체적으로, 상기 대장균 속 (Escherichia sp.) 미생물은 대장균 (Escherichia coli) 일 수 있다.
구체적으로, 상기 효모 (yeast)는 사카로미세스 세레비시아 (Saccharomyces cerevisiae), 또는 캔디다 유틸리스 (Candida utilis)인 것일 수 있다.
본 발명의 일 예에 따른 재조합 미생물은 푸코오스 합성능을 가지거나, 푸코오스 합성능이 개선된 것일 수 있다. 2'-푸코실락토오스를 생산하기 위해 어떤 미생물이든 기본적으로 외래의 α-1,2-푸코오스 전이효소를 도입해야 하는 것은 동일하나, 각 미생물의 균주 자체의 유전적 특성이 다르기 때문에, 대장균이나 바실러스 메가테리움은 α-1,2-푸코오스 전이 효소만 삽입해도 2'-푸코실락토오스 생산이 가능하나, 바실러스 써틸리스, 코리네박테리움, 효모 미생물 등은 추가적으로 푸코오스 합성 효소를 도입할 수 있다.
구체적으로, 본 발명의 일 예에 따른 재조합 미생물은 GDP-D-만노오스-4,6-데하이드라타아제 (GDP-D-mannose-4,6-dehydratase), GDP-L-푸코오스 신타아제 (GDP-L-fucose synthase, WcaG), 포스포만노뮤타아제 (phosphomannomurase), 및 GTP 만노오스-1-포스페이트 구아닐릴트랜스퍼라아제 (GTP-mannose-1-phosphate guanylyltransferase)로 이루어지는 군에서 선택된 1종 이상을 보유하거나, 과발현하는 것을 특징으로 할 수 있다. 이에, 본 발명의 일 예에 따른 재조합 미생물은, α-1,2-푸코오스 전이효소 발현 특성과 더불어, 푸코오스 합성능이 개선되어 α-1,2-푸코오스 생산 수율이 더욱 향상될 수 있다.
예를 들어, 상기 푸코오스 합성 효소는 서열번호 19 내지 22로 이루어지는 군에서 선택된 1종 이상의 아미노산 서열을 포함하는 것일 수 있다.
예를 들어, 상기 GTP 만노오스-1-포스페이트 구아닐릴트랜스퍼라아제는 서열번호 19의 아미노산 서열을 포함하는 것이거나, 서열번호 23의 핵산 서열에 의해 암호화되는 것일 수 있다.
예를 들어, 상기 포스포만노뮤타아제는 서열번호 20의 아미노산 서열을 포함하는 것이거나, 서열번호 24의 핵산 서열에 의해 암호화되는 것일 수 있다.
예를 들어, GDP-L-푸코오스 신타아제 (WcaG)는 서열번호 21의 아미노산 서열을 포함하는 것이거나, 서열번호 25의 핵산 서열에 의해 암호화되는 것일 수 있다.
예를 들어, 상기 GDP-D-만노오스-4,6-데하이드라타아제 (GDP-D-mannose-4,6-dehydratase)는 서열번호 22의 아미노산 서열을 포함하는 것이거나, 서열번호 26의 핵산 서열에 의해 암호화되는 것일 수 있다.
본 발명의 일 예에 따른 재조합 미생물은 락토오스 막 수송 단백질을 발현 또는 과발현하는 것일 수 있다. 상기 락토오스 막 수송 단백질은 2’-푸코실락토오스의 기질인 락토오스를 세포 내로 도입하여, 2’-푸코실락토오스 생산능을 개선하는 것일 수 있다.
구체적으로, 상기 락토오스 막 수송 단백질은 Lac12 및/또는 LacY인 것일 수 있다.
예를 들어, 상기 Lac12는 서열번호 27의 아미노산 서열을 가지는 것일 수 있다. 예를 들어, 상기 Lac12를 암호화하는 핵산은 서열번호 29의 핵산 서열을 포함하는 것일 수 있다. 일 예로, 상기 Lac12는 서열번호 29의 핵산 서열에 의해 암호화되는 것일 수 있다.
예를 들어, 상기 LacY는 서열번호 28의 아미노산 서열을 가지는 것일 수 있다. 예를 들어, 상기 LacY를 암호화하는 핵산 서열은 서열번호 30의 핵산 서열을 포함하는 것일 수 있다. 일 예로, 상기 LacY는 서열번호 30의 핵산 서열에 의해 암호화되는 것일 수 있다.
본 발명의 또 다른 일 예는, 본 발명에 따른 재조합 미생물을 이용하여 얻어진 α-1,2-푸코오스 전이효소, 상기 미생물의 균체, 상기 미생물의 배양물, 상기 미생물의 파쇄물, 및 상기 파쇄물 또는 배양물의 추출물로 이루어진 군에서 선택된 1종 이상을 포함하는, 2'-푸코실락토오스 생산용 조성물에 관한 것이다. 상기 재조합 미생물, 상기 α-1,2-푸코오스 전이효소, 상기 2'-푸코실락토오스 등은 전술한 바와 같다.
상기 배양물은 상기 재조합 미생물로부터 생산된 효소를 포함하는 것으로, 상기 재조합 미생물을 포함하거나, 상기 재조합 미생물을 포함하지 않는 무세포(cell-free) 형태일 수 있다. 상기 파쇄물은 상기 재조합 미생물을 파쇄한 파쇄물 또는 상기 파쇄물을 원심분리하여 얻어진 상등액을 의미하는 것으로, 상기 재조합 미생물로부터 생산된 효소를 포함하는 것이다. 본 명세서에 있어서, 별도의 언급이 없는 한, 2'-푸코실락토오스 제조에 사용되는 재조합 미생물은 상기 미생물의 균체, 상기 미생물의 배양물 및 상기 미생물의 파쇄물로 이루어진 군에서 선택된 1종 이상을 의미하는 것으로 사용된다.
본 발명의 또 다른 일 예는, 락토오스 (lactose)를 포함하는 배지에서 본 발명에 따른 재조합 미생물을 배양하는 단계를 포함하는, 2’-푸코실락토오스 (2’-fucolsylactose)의 생산방법에 관한 것이다.
상기 제조방법은 배양물에서 얻어진 α-1,2-푸코오스 전이효소, 상기 미생물의 균체, 상기 미생물의 배양물, 상기 미생물의 파쇄물, 및 상기 파쇄물 또는 배양물의 추출물로 이루어진 군에서 선택된 1종 이상을 락토오스 함유 원료와 반응하여 2'-푸코실락토오스를 생산하는 것일 수 있다.
본 발명은 바이오 세이프 레벨 1 (biosafety level 1) 균주에서 유래한 푸코오스 전이효소가 도입되어, 안정성 측면에서 소비자의 우려가 없는 재조합 미생물 및 이를 이용한 2’-푸코실락토오스의 생산방법을 제공할 수 있다.
도 1은 본 발명의 실시예에 따라 선별된 효소들의 단백질 발현 SDS-PAGE 결과 및 예상 단백질 크기를 나타낸 도면이다.
도 2는 본 발명의 실시예에 따른 플라스크 반응 (상단) 및 시험관 반응 (하단)의 2FL 생산 경향성을 나타낸 도면이다.
도 3은 본 발명의 일 예에 따른 α-1,2-푸코오스 전이효소의 E.coli host를 이용한 2FL 생산 활성 비교 결과를 나타낸 도면이다.
도 4는 B.megaterium 14581 host를 이용한 다양한 2FL생산성 비교 결과를 나타낸 도면이다.
도 5는 본 발명의 실시예에 따른 GDP-fucose cassette를 나타낸 도면이다.
도 6은 본 발명의 실시예에 따른 GDP-fucose cassette를 도입한 균주를 이용하여 2FL 생산성을 확인한 도면이다.
도 2는 본 발명의 실시예에 따른 플라스크 반응 (상단) 및 시험관 반응 (하단)의 2FL 생산 경향성을 나타낸 도면이다.
도 3은 본 발명의 일 예에 따른 α-1,2-푸코오스 전이효소의 E.coli host를 이용한 2FL 생산 활성 비교 결과를 나타낸 도면이다.
도 4는 B.megaterium 14581 host를 이용한 다양한 2FL생산성 비교 결과를 나타낸 도면이다.
도 5는 본 발명의 실시예에 따른 GDP-fucose cassette를 나타낸 도면이다.
도 6은 본 발명의 실시예에 따른 GDP-fucose cassette를 도입한 균주를 이용하여 2FL 생산성을 확인한 도면이다.
이하, 본 발명을 하기의 실시예에 의하여 더욱 상세히 설명한다. 그러나 이들 실시예는 본 발명을 예시하기 위한 것일 뿐이며, 본 발명의 범위가 이들 실시예에 의하여 한정되는 것은 아니다.
실시예 1. 생물 정보학 도구를 이용한 a-1,2-푸코오스 전이효소 스크리닝
생물 정보학 도구를 이용하여 Biosafety level 1 유래 a-1,2-fucosyltransferase 효소 (이하, 2FT) 후보를 선별하였다. 구체적으로, ncbi (https://www.ncbi.nlm.nih.gov/protein/)에서 a-1,2-푸코오스 전이효소 기능을 가질 것으로 예상되는 효소 중 Bacteria 유래 및 GRAS bacteria 유래를 기준으로 >90% 상동성 sequence를 제거하고, 단백질 형태 시각화 프로그램 (http://wlab.ethz.ch/protter)을 이용하여 막 단백질의 유무를 파악하였다.
컴퓨터 생물 정보학 도구들을 사용하여 32개의 후보군을 선별하였다 (표 1). 또한 단백질 발현에 문제가 되는 막단백질의 존재 여부를 프로그램을 통하여 예측 하였을 때에 32개 후보군 중, Lactococcus lactis 유래를 제외한 나머지 31개의 효소에서 극명한 막단백질이 예측되지 않았다.
선별 기준 | 후보군 |
Bacteria | 6431 |
GRAS bacteria (ex) Acetobacter, Bacillus, Lactobacillus etc) |
67 |
중복 제거 (>90% 상동성 sequence 제거,partial swquence 제거) |
32 |
후보군 중 Biosafety level 1 Bm2FT가 반응성이 이후 확인되어, Bm2FT 단백질 서열 상동성을 바탕으로 추가 후보군을 Bacillus megaterium 유래 효소를 탐색하였다. Bm2FT의 서열 정보를 바탕으로 BLAST 프로그램 (https://blast.ncbi.nlm.nih.gov/Blast.cgi)을 이용하여 효소를 탐색하였다. 선별된 효소 후보군 목록을 표 2에 나타내었다.
명명 | 서열 | 서열번호 |
Am2FT_2 | MNMERKTGLMNKKYVSPCFLPGMRLGNIMFTLAAACAHARTVGVECRVPWAYNDASLMLRSRLGGWVLPSTPCGTNEPPSWQEPSFAYCPVPSRIRTGGLRGYFQSARYFEGQEAFIRALFAPLTAEKEPGAVGIHIRLGDYRRLRDKHRILDPGFLRRAAGHLSSGKNRLVLFSDEPDEAAEMLARVPAFGRFALEIDRGAPCESLRRMTAMEELVMSCSSFSWWGAWLGNTRKVIVPRDWFVGGVEDYRDIYLPHWVTL | 1 |
Bm2FT | MKIVQISSGLGNQLFQYALYKRLSMNNNDVFLDVETSYQLNKNQHNGYEIERIFSIQPSHATKGMIDELADVDNRLINRLRRKLFGPKNSMYTETKEFSYDCEVFTKDGIYIKGYWQNYNYFKEIEDDLKNELVFKKALDLKNSHLINQMNKEISVSIHVRRGDYYLNKEYENKFGNIADLDYYLKAINFIKKEVDDPKFYVFSDDIKWAKENLNLTDNVTYVEHNKGSDSYKDMRLMTCCKHNIIANSTFSWWGAFLNENKNKIVIAPGKWINVEGVGGINLFPEGWIVY | 2 |
Bm2FT_2 | MGCIKRLFLYEYGGRCFLVIVKIKGGFGNQLFTYASAYAISQELQQNLIMDKVIYDLDYFRKFELPSLSLKYDQMLISKFVPNTKVKTLIYKVLRRLKLKGFTEVHEKKEFSFDENIYNLSGDIYLDGYWQNYRYFHKYYKDLSEMFVPRETPRKEVTDYITSLRGVNSVAMHVRRGDYKTFNGGKCLSLDYYIKAMEYFNSNDVQFYVFTDDIDFCEKNLPNSENINYVSRSEKLTDIEEFFIMKECKNFIIANSSFSWWAAYLSEQKADSLIVAPVVDMWKRDFYPDEWVALNTHLE | 3 |
Bm2FT_3 | MIIVQIKGGLGNQLFSYASAYGIARENETELIIDRYIYDTSYSLRKYMLDFFPEIDEALLLKYIPKKNKISQIMYKLIRKSKLKYKYKAQLFLEEEEFKFTRISTQNENLYLNGYWQSYVYFDKYRNDIIKKFTPLVSFNQDGNQLLNEIKSYNSVAIHVRRGDYINFKGGKCLDSSYYIKAMKHLYNLKGKNLFFYIFTDDVEYCKRIFKNVANVKFIGEEAKLSDFEEFTLMTHCKNLIIGNSSFSWWAAYLASCKDKAVIAPVVDMWTEDFYLPEWIKIKADLQ | 4 |
Bs2FT | MKIVHISSGLGNQMFQYALYKKLSLIQDNVFLDTITSYQLYPNQHNGYELEKVFTIKPRHASKELTYNLSDLDNSVTSRIRRKLIGSKKSMYIEHKEFEYDPNLFYQENIYIKGYWQNYDYFKDIENELKNDFTFQRALDEKNNNLAIKINNENSISIHVRRGDYYLNRKNQEKFGDIANLEYYSKAISYIKERIDNPKFYIFSDNVEWVKQNLNSLEEAVYIDYNVGNDSYKDMQLMSLCKHNIIANSSFSWWGAFLNKNMEKIVIAPGKWINMKGVKKVNLFPKDWIIY | 5 |
Bm2FT_4 | MIIVRVIGGLGNQMFQYALYKSLENEGKEVKLDLTGFGDYDLHNGYELNKIFNINENVATKDEINKLIKLPDNKVLSMIKRKFFSSTINYYSQDQFKYLSEIFQLDNVYLDGYWQSEKYFQGIKEVIRKEFKFKGEPNPKNIEMVKLMRGSNSVSIHFRRGDYISNPDAYKVHGGITTIHYYENAVKEIKSKVKEPKFFIFSDDIKWVKENFKLEDAFFIDWNTGSESYRDIELMSNCKHNIIANSTFSWWGAWLNKNKNKIVIAPNQWFNTIDTEDVIPDTWQCINTF | 6 |
Bt2FT | MNEIHVLLTGRLGNQLFQYAFARSLQKQYGGRIVCNTFDLDHRSEKSKVADQKFSYAMGDFKLDENVALEDAALPWYADFSSPLIKPVKKLFPRRYFDFMARRGYLMWQRTDYMPIPKLECEDVFAWGWWQDIRYFQDVQTELSDEVVPVTDPLPENQYIYNAASGEESVCISIRCGNYFNPTVKKLLYVCTPEYFRNAVESITKRLTHPKFIVFTDDVDWVKENIRFESAYPQYEFLYERGSDTVEEKIRMMTLCKHFIISNSTFSWWAQFLSKSENKIVIAPDRWFVDGRRIGLYMHGWTLIPAGQ | 7 |
Fp2FT | MIYVEMHGRLGNQMFQYAAARALQEKNNQPIMLSFRKVIGANTEGTAGWENSLKYFNVKPCEYYMGKKSLVTEYPVEYRLLCYAYALSYKPLMNNMNRWYEYQVKCCRFLDRFGIRWIANGYYDFHYNGLKNYLLNGSFESPKYFDSIRDKLLEEFTPREEERKENKRLYEQIRKRNSVCLSVRHFQLTGKQADMYDVCSLEYYQTAIRKMCELIENPLFVVFSDDIEWVKNTIDLSRVEVVYETPGNPVWEKLRLMYSCKNFIIPNSTFAWWAQYLSRNPDKYVLCPAKWFNNNFESPLIASQWVRIDREGNIVNE | 8 |
Gs2FT | MKIVKVIGGLGNQMFQYAFYRNLKAKFQEVKLDITAFETYKLHNGYELERVFDIKPEYATKKEIYPLTTNRNSKISKIKRRIFGGKETEYIEKDLKFDPEVFKVTGDVYFEGYWQTEKYFKEIEDLIRKDFQFKNPLTNKNLELSNKIKNENSVSIHVRRGDYYTSKKAERKHGNIATIEYYQKAVRKITEFVDNPVFYIFSDDIPWVKENLKLENEVIYVDWNKGLDSYIDMQLMSICKHNIIANSTFSWWGAWLNQNKNKIVIAPSRWINNKRLDTSDVIPKEWIKI | 9 |
Lg2FT | MLYVEMDGRCGNQLFHYAVARYIQLAIGNKEKLCLNFNKIFEKKDENNGWVDYLKDFKTVPYSYYSKSGTILKNESNFIQKIAIGLKAIQIKSLTKKSRQEQADKAEVGQRTLNKLGVYWVREGVNQIYPYKNNKILVSGICESNFIYEIQEQLQKELIPVTPVSSLNKSLLEKIDNCNSVCISVRRGDFFNNKNAKKYGVCSPEYYIRAKKYFDKKRLENTVYFCFSDDIEWCKENLKFTDKNVIFVSQEMPVYETLRLMSHCKHFILSNSTFSWWGQFLSEYKDKIVVSPARWNNDGYDTNLIDKNWILIDA | 10 |
Ll2FT | MIYVEIRGNLGNQLFIYATAKKIQKLTGQKIQLNTTTLNKYFPNYKFGLSEFIMEDPDCFIESYKKLPWFTNEYLLPIKIFKKILNKTPKINKILSDFFFKAFEKKGYFIWRGETFKKFSLGNHKNYYLSGFWQSEDYFYDIRDELLEIITPINSIRECNFELLNLIRNSESICVSIRRGDYVDNPKISAIYNVCDINYFIESVNEIKKNVVNVKVICFSDDVEWVKKNIKFDCETHYETYGNSLSEKVQLMSSCKHFVLSNSSFSWWTEFLSIRGGITIAPKNWYADEREADIYRKNWIYLEDKTEEE | 11 |
Ls2FT | MEEIHTLLTGRLGNQLFQYAFARNLQKQYGGQIYCDVYELEHRMSKVADEKFSYAMSGFKLDAGVIREDQAFPWYADFSNPVIKPIKKAMPRKYFELMARRGYLMWQRSDYMPIPVLDTQRVFASGWWQDIRYLQNVQEELSDEIVPITNPLQENRYIYDAAHDRDSVCISIRCGNYFNPTVKKLLYVCNPQYFHDSVKRISQMLAHPKFIVFTDDVSWVKEHLKFEDTYPQFEFLYERGCDTAEEKIRMMAMCNNFIISNSTFSWWAQFLSKNKEKIVIAPDKWFVDGRKIGLYMDGWTLVPAGR | 12 |
Ri2FT | MIAVKIGDGMGNQLFNYACGYAQARRDGDSLVLDISECDNSTLRDFELDKFHLKYDKKESFPNRNLGQKIYKNLRRALKYHVIKEREVYHNRDHRYDVNDIDPRVYKKKGLRNKYLYGYWQHLAYFEDYLNEITAMMTPAYEQSETVKKLQEEFKKTPTCAVHVRGGDIMGPAGAYFKHAMERMEQEKPGVRYIVFTNDMERAEEALAPVLESQKKDAVGQAENRLEFVSEMGEFSDVDEFFLMAACQNQILSNSTFSTWAAYLNQNPDKTVIMPDDLLSERMRQKNWIILK | 13 |
비교예 1. 종래의 α-1,2-푸코오스 전이효소의 준비
helicobactor pyori 유래의 2FT (FutC)는 2FT 효소 중 활성이 가장 높아 여러 문헌에서 2FL 생산을 위해 사용하고 있으므로 대조군으로 선정하였다.
Microbiol Res. 2019 May;222:35-42 논문 Search for bacterial α1,2-fucosyltransferases for whole-cell biosynthesis of 2'-fucosyllactose in recombinant Escherichia coli.에서 대장균을 이용하여 Thermosynechococcus elongates 유래의 2FT 반응성을 확인하였으며, 이를 대조군으로 선정하였다.
WO 2015/175801 문헌에서 Akkermansia muciniphila 유래의 2FT 반응성을 확인하지 못하였지만, 생산성을 재검토 차 대조군으로 선정하였다.
선별된 대조군 효소 목록 및 정보를 표 3에 나타냈다.
이름 | 공여체 | accession # |
Te2FT | Thermosynechococcus elongatus | WP_011056838.1 |
Fut C (Hp2FT) |
helicobactor pyori | WP_080473865.1 |
Am2FT_1 | Akkermansia muciniphila | WP_081429121.1 |
실시예 2. 대장균용 발현 플라스미드 클로닝
실시예 1에서 선별된 효소와 비교예 1의 대조군 효소의 대장균에서의 활성을 확인하기 위해 효소 발현 플라스미드에 클로닝을 진행하였다.
구체적으로, 실시예 1에서 선별된 효소 서열의 코돈을 대장균에 최적화하고자 코돈 최적화 프로그램 (http://genomes.urv.es/OPTIMIZER/)에 서열 입력 후, Escherichia_coli 를 선택하여 guided random 코돈 최적화 검색을 수행하였다. 확인된 효소 서열을 합성하였고, 조건에 맞추어 표 5의 제시된 프라이머을 통해 PCR를 진행하여 합성 유전자를 증폭하였다. pET24ma의 플라스미드 증폭의 경우, 합성 유전자의 stop codon의 서열이 TAG 혹은 TAA인지에 맞추어 TAG_PET24ma_F 혹은 TAA_PET24ma_F PCR 프라이머를 경우에 따라 사용하였다. 각 효소에 따른 프라이머 서열 정보 및 이름을 표 4에 나타냈다.
명명 | 프라이머 서열 (5’→ 3’) | 서열번호 |
Te2FT_F | catatgATTATCGTTCATCTGTG | 31 |
Te2FT_R | CTCGAGTCACAGAACAATCCACC | 32 |
Bt2FT_F | catatgAACGAAATCCACGTGCT | 33 |
Bt2FT_R | CTCGAGTCACTGGCCAGCCGGAATC | 34 |
Bm2FT_F | catatgAAAATTGTTCAGATTTCT | 35 |
Bm2FT_R | CTCGAGTCAGTAAACGATCCAACCT | 36 |
Lg2FT_F | catatgCTGTACGTTGAAATGGACG | 37 |
Lg2FT_R | CTCGAGTTACGCGTCGATCAGGATC | 38 |
Ls2FT_F | catatgGAAGAGATTCACACCCTG | 39 |
Ls2FT_R | CTCGAGCTAACGGCCGGCAGGTAC | 40 |
Ll2FT_F | catatgTCTAAAAACATTTATGTAC | 41 |
Ll2FT_R | CTCGAGTTAGATTTTGATCCAGTTGT | 42 |
Am2FT_1_F | aagaaggagatatacatatgATGGCTGGACATTCTTGCG | 43 |
Am2FT_1_R | tggtggtggtggtgctcgagTTAAATGCGCAGCCATGAGC | 44 |
Am2FT_2_F | aagaaggagatatacatatgATGAACATGGAACGTAAGA | 45 |
Am2FT_2_R | tggtggtggtggtgctcgagTTACAGGGTCACCCAGTGCG | 46 |
Fp2FT_F | aagaaggagatatacatatgATGATTTACGTGGAAATG | 47 |
Fp2FT_R | tggtggtggtggtgctcgagTTATTCGTTAACGATGTTA | 48 |
Ri2FT_F | aagaaggagatatacatatgATGATCGCAGTTAAGATC | 49 |
Ri2FT_R | tggtggtggtggtgctcgagTTATTTCAGGATAATCCAGT | 50 |
Bm2FT_2_F | aagaaggagatatacatatgGGTTGCATTAAACGCCTGT | 51 |
Bm2FT_2_R | TGGTGGTGGTGGTGCTCGAGCTATTCCAGGTGAGTGTTCA | 52 |
Bm2FT_3_F | aagaaggagatatacatatgATCATTGTCCAGATTAAA | 53 |
Bm2FT_3_R | TGGTGGTGGTGGTGCTCGAGCTATTGAAGATCCGCTTTGAT | 54 |
Bm2FT_4_F | aagaaggagatatacatatgATCATTGTGCGTGTTATC | 55 |
Bm2FT_4_R | TGGTGGTGGTGGTGCTCGAGTTAGAACGTATTAATACACT | 56 |
Bs2FT_F | aagaaggagatatacatatgAAAATTGTAAAAGTTATC | 57 |
Bs2FT_R | TGGTGGTGGTGGTGCTCGAGTTAGTAGATGATCCAGTCTT | 58 |
Gs2FT_F | aagaaggagatatacatatgAAAATTGTAAAAGTTA | 59 |
Gs2FT_R | TGGTGGTGGTGGTGCTCGAGTTAGATTTTAATCCATTCT | 60 |
TAG_PET24ma_F | TAGctcgagcaccaccaccaccacc | 61 |
TAA_PET24ma_F | TAActcgagcaccaccaccaccacc | 62 |
PET24ma_R | catatgtatatctccttcttaaagttaaacaaaattattt | 63 |
Te2FT, Bt2FT, Bm2FT, Lg2FT, Ls2FT, 및 Ll2FT 합성 유전자의 경우 유전자 정제 후, 각각 1uL NEB 제한효소 (Nde1/ Xho1)을 처리하여 37℃에서 3시간 보관하였다. 이후 제한 효소와 버퍼를 제거하기 위해 다시 유전자 정제 후 16℃에서 동일 제한 효소를 처리한 pET24ma 와 함께 4시간 ligation을 진행하였다. 상기 Te2FT, Bt2FT, Bm2FT, Lg2FT, Ls2FT, 및 Ll2FT 합성 유전자를 제외한 나머지 합성 유전자는 모두 Gibson assembly 방법으로 cloning을 진행하기 위해 gel prep 을 하였다.
Prep 후 nano drop으로 각각의 유전자 농도를 확인하고, vector와 insert의 비가 1:2가 되며 총 반응부피가 8uL이 되도록 2X NEBuilder® HiFi DNA Assembly Master Mix를 넣어주었다. 이후 50℃에서 1시간 동안 반응하였다. cloning 유전자를 DH10b E.coli competent cell에 transformation 한 후, Km 항생제가 들어간 plate에 도말하여 overnight으로 키웠다.
각각의 plate에 자란 colony중 5개씩을 선별하여 유전자에 해당하는 표 5 프라이머를 이용하여 colony PCR을 수행하였다. DNA gel 상에 크기에 해당하는 band를 확인한 colony를 Km 항생제가 포함된 3ml LB 배지에 접종 후, overnight 키웠다. 다음날, cell을 모아 DNA를 추출하고 스퀀싱 하였다.
실시예 3. 효소 단백질 발현 확인
대장균 내에서 cloning한 각각의 유전자가 단백질 발현하고 있는지 확인하였다. E.coli BL21(DE3) △fucl△fucK::AprR lacZ△m15 ::PlacUV5 competent cell에 실험할 각각의 2FT vector들과 GDP-fuc를 생산하는 효소를 지닌 pCDFm::PT7manC-manB와 PT7-gmd-wcaG 를 함께 transformation 하였다. Km와 Amp 항생제가 들어간 plate에 도말하여 selection 하였다.
colony를 Km와 Amp 항생제가 포함된 3mL 배지에 접종 후, seed를 overnight 키웠다. 4mL LB에 200uL seed와 0.1mM IPTG를 첨가 후 30℃에서 16시간 키웠다. 4mL에서 자란 세포를 원심분리기로 모두 수거한 후 상등액을 버렸다. 500uL의 버퍼를 첨가 후 bead beater로 파쇄 후 13000 rpm에서 20분간 원심 분리하여 soluble 단백질을 수거하였다. size marker와 함께 SDS-PAGE 젤을 내렸다. 선별 효소들의 total 및 soluble 단백질 발현 SDS-PAGE 결과 (도 1의 좌측) 및 예상 단백질 크기 (도 1의 우측)를 도 1에 나타냈다. 도 1에서 ManB, ManC, Gmd, 및 WcaG는 추가적으로 넣어준 pCDFm::PT7manC-manB + PT7-gmd-wcaG 플라스미드에서 발현된 단백질이다.
도 1에 나타난 바와 같이, SDS-PAGE를 통하여 비교한 단백질 중 8개의 효소 Bm2FT, Te2FT, Hp2FT, Am2FT_2, Fp2FT, Ri2FT, Bm2FT_2, 및 Bm2FT_3에서 과발현을 확인하였다.
실시예 4. 시험관을 이용한 반응 분석 최적화
250mL 플라스크에서 총 50 mL culture를 통해 2FL 생산성을 확인하였다. 효소 후보군이 많은 경우 확실한 비교분석을 위해 3반복 이상의 실험 재연이 필수적이나, 기존 50 mL 배양 조건의 경우 분석시 필요량 (1mL) 이상의 노동력이 드는 단점이 있다. 다양한 후보군의 빠른 반응 경향성 파악을 위해 10mL 시험관에서 4mL 배양을 통해 효소 반응 확인 방법을 확립하고자 한다.
Hp2FT, Bm2FT, Bt2FT 가 클로닝된 plasmid를 각각 pCDFm::PT7manC-manB + PT7-gmd-wcaG plasmid와 함께 E.coli 균주에 transformation 하였다. 각각의 plate 에서 3개의 colony를 접종하여 overnight seed culture를 진행하였다.
seed 중 500uL는 50mL flask culture 에 induction 하였다. OD600 이 ~0.8 정도가 될 때 0.1mM IPTG와 10mM lactose를 넣어 준 후 20h 후 HPLC로 분석하였다.
test tube 반응의 경우, seed 200 uL를 0.1mM IPTG와 10mM lactose가 첨가된 4mL 배지에 첨가하였다. test tube 반응은 flask의 OD를 확인하는 step을 생략할 수 있다.
20h 후 HPLC 로 분석하여 플라스크 반응 (도 2의 상단) 및 시험관 반응 (도 2의 하단)의 2FL 생산 경향성을 도 2에 나타냈다. 도 2에 나타난 바와 같이, 정립한 시험관 반응 조건이 플라스크 조건보다는 2FL의 생산적이 1/3 수준이나, 그 반응 경향성은 동일하여 다양한 효소를 한번에 비교 분석하는 것에 적합함을 확인하였다.
실시예 5. HPLC 분석을 통한 2FL 생산성 비교
실시예 1에서 선별된 효소 및 대조군 효소 3종의 반응성을 비교하였다.
비교할 플라스미드를 각각 E.coli 에 transformation 후, 37℃에서 키웠다. 각각의 plate 에서 3개의 colony를 접종하여 overnight seed culture를 진행하였다. seed 200 uL를 0.1mM IPTG와 10mM lactose가 첨가된 4mL 배지에 첨가하였다. 20h 후 HPLC 로 분석하였다. E.coli host를 이용한 다양한 2FT의 2FL 생산 활성 비교 결과를 도 3에 나타냈다.
도 3에 나타난 바와 같이, FutC 대비 동등 또는 높은 활성 효소 3개를 포함하여, 2FL 생산 효소 총 5개를 확보하였다: Am2FT_2, Bm2FT, Bm2FT_2, Bm2FT_3, 및 Bs2FT 의 경우 FutC 대비 각각 122%, 207%, 84%, 100%, 21% 2FL 생산 양상을 HPLC 분석을 통해 확인하였다. 2FL 생산 효소 5종의 서열을 표 5에 나타냈다.
명명 | 서열 | 서열번호 |
Am2FT_2 | MNMERKTGLMNKKYVSPCFLPGMRLGNIMFTLAAACAHARTVGVECRVPWAYNDASLMLRSRLGGWVLPSTPCGTNEPPSWQEPSFAYCPVPSRIRTGGLRGYFQSARYFEGQEAFIRALFAPLTAEKEPGAVGIHIRLGDYRRLRDKHRILDPGFLRRAAGHLSSGKNRLVLFSDEPDEAAEMLARVPAFGRFALEIDRGAPCESLRRMTAMEELVMSCSSFSWWGAWLGNTRKVIVPRDWFVGGVEDYRDIYLPHWVTL | 1 |
Bm2FT | MKIVQISSGLGNQLFQYALYKRLSMNNNDVFLDVETSYQLNKNQHNGYEIERIFSIQPSHATKGMIDELADVDNRLINRLRRKLFGPKNSMYTETKEFSYDCEVFTKDGIYIKGYWQNYNYFKEIEDDLKNELVFKKALDLKNSHLINQMNKEISVSIHVRRGDYYLNKEYENKFGNIADLDYYLKAINFIKKEVDDPKFYVFSDDIKWAKENLNLTDNVTYVEHNKGSDSYKDMRLMTCCKHNIIANSTFSWWGAFLNENKNKIVIAPGKWINVEGVGGINLFPEGWIVY | 2 |
Bm2FT_2 | MGCIKRLFLYEYGGRCFLVIVKIKGGFGNQLFTYASAYAISQELQQNLIMDKVIYDLDYFRKFELPSLSLKYDQMLISKFVPNTKVKTLIYKVLRRLKLKGFTEVHEKKEFSFDENIYNLSGDIYLDGYWQNYRYFHKYYKDLSEMFVPRETPRKEVTDYITSLRGVNSVAMHVRRGDYKTFNGGKCLSLDYYIKAMEYFNSNDVQFYVFTDDIDFCEKNLPNSENINYVSRSEKLTDIEEFFIMKECKNFIIANSSFSWWAAYLSEQKADSLIVAPVVDMWKRDFYPDEWVALNTHLE | 3 |
Bm2FT_3 | MIIVQIKGGLGNQLFSYASAYGIARENETELIIDRYIYDTSYSLRKYMLDFFPEIDEALLLKYIPKKNKISQIMYKLIRKSKLKYKYKAQLFLEEEEFKFTRISTQNENLYLNGYWQSYVYFDKYRNDIIKKFTPLVSFNQDGNQLLNEIKSYNSVAIHVRRGDYINFKGGKCLDSSYYIKAMKHLYNLKGKNLFFYIFTDDVEYCKRIFKNVANVKFIGEEAKLSDFEEFTLMTHCKNLIIGNSSFSWWAAYLASCKDKAVIAPVVDMWTEDFYLPEWIKIKADLQ | 4 |
Bs2FT | MKIVHISSGLGNQMFQYALYKKLSLIQDNVFLDTITSYQLYPNQHNGYELEKVFTIKPRHASKELTYNLSDLDNSVTSRIRRKLIGSKKSMYIEHKEFEYDPNLFYQENIYIKGYWQNYDYFKDIENELKNDFTFQRALDEKNNNLAIKINNENSISIHVRRGDYYLNRKNQEKFGDIANLEYYSKAISYIKERIDNPKFYIFSDNVEWVKQNLNSLEEAVYIDYNVGNDSYKDMQLMSLCKHNIIANSSFSWWGAFLNKNMEKIVIAPGKWINMKGVKKVNLFPKDWIIY | 5 |
실시예 6. Corynebacterium 균주 내 푸코실 전이 효소를 이용한 2’-FL 생산성 비교
플라스미드 제작 및 2'-푸코실락토오스 (fucosyllactose, 2'-FL)의 생산을 위해 코리네박테리움 글루타미쿰 (C. glutamicum) ATCC 13032를 이용하였다. pMBC 플라스미드를 구축하기 위해 코리네박테리움 글루타미쿰 ATCC 13032의 genomic DNA로부터 두 개의 DNA primer를 이용한 PCR 반응을 통해 manB 유전자를 증폭한 후, pCES208 플라스미드에 이를 삽입시켰다. 구축된 상기 플라스미드에 다시 코리네박테리움 글루타미쿰 ATCC 13032의 genomic DNA로부터 두 개의 DNA primer를 이용한 PCR 반응을 통해 manC 유전자를 증폭하였고, 플라스미드에 이를 후 삽입시킴으로써 pMBC를 구축하였다. 또한, pEGW 플라스미드를 구축하기 위해 대장균인 K-12 의 genomic DNA로부터 두 개의 DNA primer 를 이용한 PCR반응을 통해 gmd-wcaG 유전자 클러스터를 증폭한 후, 플라스미드에 삽입시킴으로써 pEGW를 구축하였다. 또한, 서열번호 2의 푸코실 전이 효소를 도입하였다.
C.glutamicum host를 이용시, Bm2FT와 GDP-fucose cassette를 도입한 균주를 10 mM lactose가 첨가된 4 mL 배지에 첨가하여 2FL을 생산하였으며, 배양 60시간이 지나면 약 120 mg/L의 2FL이 생산되었다. pMBC와 pEGW 공벡터를 넣어준 균주의 경우 2FL를 생산하지 않았다.
실시예 8. Bacillus megaterium 균주 내 푸코실 전이 효소를 통한 2’-FL 생산성 비교
E.coli host 사용시 FutC 대비 가장 더 많은 활성이 증대된 Am2FT_2, Bm2FT 효소를 각각 B. megaterium 14581 (한국 미생물 보존 센터 (KCCM) 기탁번호 40441) host 내에 형질전환 하여 각각의 2FL 생산성을 비교하였다. 반응성의 정확한 확인을 위해 대조군으로는 2FT 유전자가 들어가지 않은 공벡터 p1525가 삽입된 b. megaterium 14581를 사용하였다. 도 4는 B.megaterium 14581 host를 이용한 다양한 2FL생산성 비교 결과를 나타낸 도면이다.
도 4에 나타난 바와 같이, B. megaterium host를 이용시, p1525 공백터를 넣어준 균주의 경우 2FL를 생산 하지 않았으며, Bm2FT와 FutC 경우, 반응 64시간이 지나면 각각 평균 40.3 mg/L 및 18.8 mg/L의 2FL이 생산 되었다. 이를 통해 Bm2FT 효소가 FutC 대비 2.1배 이상의 생산성 증대를 보임을 확인하였다.
B. megaterium 균주를 사용하여서 모유 올리고당을 생산한 연구가 현재까지 보고된 바 없으며, B. megaterium 14581 host는 biosafety level 1 균주로 인체에 무해하므로, 본 발명에서 확립된 Bm2FT를 GRAS 균주에 도입하여 기존 효소 (FucT) 대비 2배 이상의 생산성을 확립하였다.
실시예 9. Bacillus subtilis 균주 내 푸코실 전이 효소를 통한 2’-FL 생산성 비교
E.coli host 사용시 FutC 대비 가장 많은 활성이 증대한 Bm2FT 효소를 Bacillus subtilis host 내에 형질전환 하여 각각의 2’-FL 생산성을 비교하였다. 반응성의 정확한 확인을 위해 대조군으로 2’-FT 유전자가 클로닝되지 않은 공벡터 pBE-S 가 형질전환 된 B. subtilis 168 (기탁번호: (ATCC® 23857™))을 사용하였다. 사용한 벡터는 B.subtilis secretion용인 pBE-S이며 상시 발현이 가능한 aprE promoter가 장착되어 있다. cloning 시에는 secretion 기능이 필요 없기 때문에 2’-FT 클로닝 진행할 때 aprE signal peptide는 제거하였다.
B.subtilis 168은 GDP-fucose를 생산하는 4종의 효소를 갖고 있지 않기 때문에, 하나의 vector에 4종의 효소를 encoding하는 유전자를 한번에 클로닝하였다. 4종 유전자 (manB, manC, gmd, wcaG)는 E.coli 유래로 PCR을 통해 유전자 증폭하여 확보하였다. 4종의 유전자를 한번에 클로닝하기 위해 사용한 vector는 p3STOP1623-2RBShp로서 E.coli-Bacillus shuttle vector이다. 본 vector는 B.megaterium 뿐만 아니라 B.subtilis에서도 사용할 수 있다. 본 Vector의 특징으로는 2개의 RBS를 갖고 있어 다중 유전자의 클로닝에 매우 용이하다. 본 vector의 promoter는 xylose inducible promoter인 PxylA이다. GDP-fucose cassette는 도 5와 같은 유전자 순서로 제작하였다.
Vector와 4종의 유전자는 앞서 Bm2FT cloning 때와 동일하게 Gibson assembly 방법을 적용하여 진행하였다. 단지 Vector의 크기가 약 6.6 Kb로 한번에 PCR하기에는 시퀀스 에러가 발생할 수 있으므로 Vector 중간에 primer를 제작하여 반으로 나누어 PCR을 진행하였다. 총 6개의 PCR fragments를 정제하기 위해 PCR mixture 전부를 전기영동하여 gel extraction 하였다. 정제하여 확보한 PCR fragments는 DNA 농도를 측정하고, 농도와 사이즈에 맞게 함량을 조절하여 Gibson assembly를 진행하였다. 그 후, 동일하게 E.coli에 형질전환 하여 single colony를 확보한 후, 콜로니 PCR을 통해 plasmid의 삽입을 확인하였다. PCR 확인된 콜로니는 LB에 배양하여 plasmid 추출 후, 시퀀스 분석을 통해 4종의 유전자가 제대로 cloning 된 것을 확인하였다. 시퀀스까지 확인된 plasmid는 생산성 확인을 위해 B.subtilis 168에 형질전환 하였다. E.coli에 형질전환하여 확인한 방법과 동일하게 콜로니 PCR로 plasmid의 삽입을 확인한 후, 2’-FL 생산성 비교를 위해 배양을 진행하였다.
도 6에 나타난 바와 같이, B.subtilis host를 이용시, pBE-S와 p3STOP1623-2RBShp 공벡터를 넣어준 균주의 경우 2FL를 생산하지 않았으며, Bm2FT와 GDP-fucose cassette를 도입한 균주의 경우, 반응 63시간이 지나면 약 150 mg/L의 2FL이 생산되었다. 이는 종래의 FutC의 평균 2FL 생산량 18.8 mg/L 대비 약 7배 이상으로 생산성이 향상된 결과이다.
실시예 10. Saccaromyces Cerevisiae 균주 내 푸코실 전이 효소를 통한 2’-FL 생산성 비교
플라스미드 제작 및 2'-푸코실락토오스 (fucosyllactose, 2'-FL)의 생산을 위해 사카로마이세스 세레비지애 (S. cerevisiae) CEN.PK2-1c를 이용하였다. 사카로마이세스 세레비지애에 없는 GDP-L-Fucose 유전자를 대장균인 K-12에서 클로닝하여 사용하였다. pRSK426 플라스미드를 구축하기 위해 대장균인 K-12 의 genomic DNA로부터 두 개의 DNA primer 를 이용한 PCR반응을 통해 gmd-wcaG 유전자 클러스터를 증폭한 후, 플라스미드에 삽입시킴으로써 pRSK426를 구축하였다. E.coli host 사용시 FutC 대비 가장 더 많은 활성이 증대된 Am2FT_2, Bm2FT 효소를 각각 S. cerevisiae host 내에 형질전환 하여 각각의 2FL 생산성을 비교하였다. 3종의 재조합 플라스미드는 사카로마이세스 세레비지애 (S. cerevisiae) CEN.PK2-1c 균주로 형질전환 후 G418 항생제가 포함된 YPD 고체 배지에서 선별하였다. 선별된 균주들은 G418 항생제가 포함된 50mL 액체 생산 배지에서 2일 배양한 뒤, 2FL 생산성을 HPLC 분석하여 측정하였다. S. cerevisiae CEN.PK2-1c host를 이용시, pRSK426 공벡터를 넣어준 균주의 경우 2FL를 생산하지 않았으며, Bm2FT, Am2FT_2와 GDP-fucose cassette를 도입한 균주의 경우, 배양 48시간이 지나면 각각 80 mg/L, 40 mg/L의 2FL이 생산되었다.
<110> SAMYANG CORPORATION
<120> Recombinant microorganism expressing fucosyltransferase and
Method of producing 2'-fucolsylactose using thereof
<130> DPP20205427KR
<160> 63
<170> koPatentIn 3.0
<210> 1
<211> 261
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Am2FT_2
<400> 1
Met Asn Met Glu Arg Lys Thr Gly Leu Met Asn Lys Lys Tyr Val Ser
1 5 10 15
Pro Cys Phe Leu Pro Gly Met Arg Leu Gly Asn Ile Met Phe Thr Leu
20 25 30
Ala Ala Ala Cys Ala His Ala Arg Thr Val Gly Val Glu Cys Arg Val
35 40 45
Pro Trp Ala Tyr Asn Asp Ala Ser Leu Met Leu Arg Ser Arg Leu Gly
50 55 60
Gly Trp Val Leu Pro Ser Thr Pro Cys Gly Thr Asn Glu Pro Pro Ser
65 70 75 80
Trp Gln Glu Pro Ser Phe Ala Tyr Cys Pro Val Pro Ser Arg Ile Arg
85 90 95
Thr Gly Gly Leu Arg Gly Tyr Phe Gln Ser Ala Arg Tyr Phe Glu Gly
100 105 110
Gln Glu Ala Phe Ile Arg Ala Leu Phe Ala Pro Leu Thr Ala Glu Lys
115 120 125
Glu Pro Gly Ala Val Gly Ile His Ile Arg Leu Gly Asp Tyr Arg Arg
130 135 140
Leu Arg Asp Lys His Arg Ile Leu Asp Pro Gly Phe Leu Arg Arg Ala
145 150 155 160
Ala Gly His Leu Ser Ser Gly Lys Asn Arg Leu Val Leu Phe Ser Asp
165 170 175
Glu Pro Asp Glu Ala Ala Glu Met Leu Ala Arg Val Pro Ala Phe Gly
180 185 190
Arg Phe Ala Leu Glu Ile Asp Arg Gly Ala Pro Cys Glu Ser Leu Arg
195 200 205
Arg Met Thr Ala Met Glu Glu Leu Val Met Ser Cys Ser Ser Phe Ser
210 215 220
Trp Trp Gly Ala Trp Leu Gly Asn Thr Arg Lys Val Ile Val Pro Arg
225 230 235 240
Asp Trp Phe Val Gly Gly Val Glu Asp Tyr Arg Asp Ile Tyr Leu Pro
245 250 255
His Trp Val Thr Leu
260
<210> 2
<211> 291
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bm2FT
<400> 2
Met Lys Ile Val Gln Ile Ser Ser Gly Leu Gly Asn Gln Leu Phe Gln
1 5 10 15
Tyr Ala Leu Tyr Lys Arg Leu Ser Met Asn Asn Asn Asp Val Phe Leu
20 25 30
Asp Val Glu Thr Ser Tyr Gln Leu Asn Lys Asn Gln His Asn Gly Tyr
35 40 45
Glu Ile Glu Arg Ile Phe Ser Ile Gln Pro Ser His Ala Thr Lys Gly
50 55 60
Met Ile Asp Glu Leu Ala Asp Val Asp Asn Arg Leu Ile Asn Arg Leu
65 70 75 80
Arg Arg Lys Leu Phe Gly Pro Lys Asn Ser Met Tyr Thr Glu Thr Lys
85 90 95
Glu Phe Ser Tyr Asp Cys Glu Val Phe Thr Lys Asp Gly Ile Tyr Ile
100 105 110
Lys Gly Tyr Trp Gln Asn Tyr Asn Tyr Phe Lys Glu Ile Glu Asp Asp
115 120 125
Leu Lys Asn Glu Leu Val Phe Lys Lys Ala Leu Asp Leu Lys Asn Ser
130 135 140
His Leu Ile Asn Gln Met Asn Lys Glu Ile Ser Val Ser Ile His Val
145 150 155 160
Arg Arg Gly Asp Tyr Tyr Leu Asn Lys Glu Tyr Glu Asn Lys Phe Gly
165 170 175
Asn Ile Ala Asp Leu Asp Tyr Tyr Leu Lys Ala Ile Asn Phe Ile Lys
180 185 190
Lys Glu Val Asp Asp Pro Lys Phe Tyr Val Phe Ser Asp Asp Ile Lys
195 200 205
Trp Ala Lys Glu Asn Leu Asn Leu Thr Asp Asn Val Thr Tyr Val Glu
210 215 220
His Asn Lys Gly Ser Asp Ser Tyr Lys Asp Met Arg Leu Met Thr Cys
225 230 235 240
Cys Lys His Asn Ile Ile Ala Asn Ser Thr Phe Ser Trp Trp Gly Ala
245 250 255
Phe Leu Asn Glu Asn Lys Asn Lys Ile Val Ile Ala Pro Gly Lys Trp
260 265 270
Ile Asn Val Glu Gly Val Gly Gly Ile Asn Leu Phe Pro Glu Gly Trp
275 280 285
Ile Val Tyr
290
<210> 3
<211> 299
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bm2FT_2
<400> 3
Met Gly Cys Ile Lys Arg Leu Phe Leu Tyr Glu Tyr Gly Gly Arg Cys
1 5 10 15
Phe Leu Val Ile Val Lys Ile Lys Gly Gly Phe Gly Asn Gln Leu Phe
20 25 30
Thr Tyr Ala Ser Ala Tyr Ala Ile Ser Gln Glu Leu Gln Gln Asn Leu
35 40 45
Ile Met Asp Lys Val Ile Tyr Asp Leu Asp Tyr Phe Arg Lys Phe Glu
50 55 60
Leu Pro Ser Leu Ser Leu Lys Tyr Asp Gln Met Leu Ile Ser Lys Phe
65 70 75 80
Val Pro Asn Thr Lys Val Lys Thr Leu Ile Tyr Lys Val Leu Arg Arg
85 90 95
Leu Lys Leu Lys Gly Phe Thr Glu Val His Glu Lys Lys Glu Phe Ser
100 105 110
Phe Asp Glu Asn Ile Tyr Asn Leu Ser Gly Asp Ile Tyr Leu Asp Gly
115 120 125
Tyr Trp Gln Asn Tyr Arg Tyr Phe His Lys Tyr Tyr Lys Asp Leu Ser
130 135 140
Glu Met Phe Val Pro Arg Glu Thr Pro Arg Lys Glu Val Thr Asp Tyr
145 150 155 160
Ile Thr Ser Leu Arg Gly Val Asn Ser Val Ala Met His Val Arg Arg
165 170 175
Gly Asp Tyr Lys Thr Phe Asn Gly Gly Lys Cys Leu Ser Leu Asp Tyr
180 185 190
Tyr Ile Lys Ala Met Glu Tyr Phe Asn Ser Asn Asp Val Gln Phe Tyr
195 200 205
Val Phe Thr Asp Asp Ile Asp Phe Cys Glu Lys Asn Leu Pro Asn Ser
210 215 220
Glu Asn Ile Asn Tyr Val Ser Arg Ser Glu Lys Leu Thr Asp Ile Glu
225 230 235 240
Glu Phe Phe Ile Met Lys Glu Cys Lys Asn Phe Ile Ile Ala Asn Ser
245 250 255
Ser Phe Ser Trp Trp Ala Ala Tyr Leu Ser Glu Gln Lys Ala Asp Ser
260 265 270
Leu Ile Val Ala Pro Val Val Asp Met Trp Lys Arg Asp Phe Tyr Pro
275 280 285
Asp Glu Trp Val Ala Leu Asn Thr His Leu Glu
290 295
<210> 4
<211> 287
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bm2FT_3
<400> 4
Met Ile Ile Val Gln Ile Lys Gly Gly Leu Gly Asn Gln Leu Phe Ser
1 5 10 15
Tyr Ala Ser Ala Tyr Gly Ile Ala Arg Glu Asn Glu Thr Glu Leu Ile
20 25 30
Ile Asp Arg Tyr Ile Tyr Asp Thr Ser Tyr Ser Leu Arg Lys Tyr Met
35 40 45
Leu Asp Phe Phe Pro Glu Ile Asp Glu Ala Leu Leu Leu Lys Tyr Ile
50 55 60
Pro Lys Lys Asn Lys Ile Ser Gln Ile Met Tyr Lys Leu Ile Arg Lys
65 70 75 80
Ser Lys Leu Lys Tyr Lys Tyr Lys Ala Gln Leu Phe Leu Glu Glu Glu
85 90 95
Glu Phe Lys Phe Thr Arg Ile Ser Thr Gln Asn Glu Asn Leu Tyr Leu
100 105 110
Asn Gly Tyr Trp Gln Ser Tyr Val Tyr Phe Asp Lys Tyr Arg Asn Asp
115 120 125
Ile Ile Lys Lys Phe Thr Pro Leu Val Ser Phe Asn Gln Asp Gly Asn
130 135 140
Gln Leu Leu Asn Glu Ile Lys Ser Tyr Asn Ser Val Ala Ile His Val
145 150 155 160
Arg Arg Gly Asp Tyr Ile Asn Phe Lys Gly Gly Lys Cys Leu Asp Ser
165 170 175
Ser Tyr Tyr Ile Lys Ala Met Lys His Leu Tyr Asn Leu Lys Gly Lys
180 185 190
Asn Leu Phe Phe Tyr Ile Phe Thr Asp Asp Val Glu Tyr Cys Lys Arg
195 200 205
Ile Phe Lys Asn Val Ala Asn Val Lys Phe Ile Gly Glu Glu Ala Lys
210 215 220
Leu Ser Asp Phe Glu Glu Phe Thr Leu Met Thr His Cys Lys Asn Leu
225 230 235 240
Ile Ile Gly Asn Ser Ser Phe Ser Trp Trp Ala Ala Tyr Leu Ala Ser
245 250 255
Cys Lys Asp Lys Ala Val Ile Ala Pro Val Val Asp Met Trp Thr Glu
260 265 270
Asp Phe Tyr Leu Pro Glu Trp Ile Lys Ile Lys Ala Asp Leu Gln
275 280 285
<210> 5
<211> 291
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bs2FT
<400> 5
Met Lys Ile Val His Ile Ser Ser Gly Leu Gly Asn Gln Met Phe Gln
1 5 10 15
Tyr Ala Leu Tyr Lys Lys Leu Ser Leu Ile Gln Asp Asn Val Phe Leu
20 25 30
Asp Thr Ile Thr Ser Tyr Gln Leu Tyr Pro Asn Gln His Asn Gly Tyr
35 40 45
Glu Leu Glu Lys Val Phe Thr Ile Lys Pro Arg His Ala Ser Lys Glu
50 55 60
Leu Thr Tyr Asn Leu Ser Asp Leu Asp Asn Ser Val Thr Ser Arg Ile
65 70 75 80
Arg Arg Lys Leu Ile Gly Ser Lys Lys Ser Met Tyr Ile Glu His Lys
85 90 95
Glu Phe Glu Tyr Asp Pro Asn Leu Phe Tyr Gln Glu Asn Ile Tyr Ile
100 105 110
Lys Gly Tyr Trp Gln Asn Tyr Asp Tyr Phe Lys Asp Ile Glu Asn Glu
115 120 125
Leu Lys Asn Asp Phe Thr Phe Gln Arg Ala Leu Asp Glu Lys Asn Asn
130 135 140
Asn Leu Ala Ile Lys Ile Asn Asn Glu Asn Ser Ile Ser Ile His Val
145 150 155 160
Arg Arg Gly Asp Tyr Tyr Leu Asn Arg Lys Asn Gln Glu Lys Phe Gly
165 170 175
Asp Ile Ala Asn Leu Glu Tyr Tyr Ser Lys Ala Ile Ser Tyr Ile Lys
180 185 190
Glu Arg Ile Asp Asn Pro Lys Phe Tyr Ile Phe Ser Asp Asn Val Glu
195 200 205
Trp Val Lys Gln Asn Leu Asn Ser Leu Glu Glu Ala Val Tyr Ile Asp
210 215 220
Tyr Asn Val Gly Asn Asp Ser Tyr Lys Asp Met Gln Leu Met Ser Leu
225 230 235 240
Cys Lys His Asn Ile Ile Ala Asn Ser Ser Phe Ser Trp Trp Gly Ala
245 250 255
Phe Leu Asn Lys Asn Met Glu Lys Ile Val Ile Ala Pro Gly Lys Trp
260 265 270
Ile Asn Met Lys Gly Val Lys Lys Val Asn Leu Phe Pro Lys Asp Trp
275 280 285
Ile Ile Tyr
290
<210> 6
<211> 289
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bm2FT_4
<400> 6
Met Ile Ile Val Arg Val Ile Gly Gly Leu Gly Asn Gln Met Phe Gln
1 5 10 15
Tyr Ala Leu Tyr Lys Ser Leu Glu Asn Glu Gly Lys Glu Val Lys Leu
20 25 30
Asp Leu Thr Gly Phe Gly Asp Tyr Asp Leu His Asn Gly Tyr Glu Leu
35 40 45
Asn Lys Ile Phe Asn Ile Asn Glu Asn Val Ala Thr Lys Asp Glu Ile
50 55 60
Asn Lys Leu Ile Lys Leu Pro Asp Asn Lys Val Leu Ser Met Ile Lys
65 70 75 80
Arg Lys Phe Phe Ser Ser Thr Ile Asn Tyr Tyr Ser Gln Asp Gln Phe
85 90 95
Lys Tyr Leu Ser Glu Ile Phe Gln Leu Asp Asn Val Tyr Leu Asp Gly
100 105 110
Tyr Trp Gln Ser Glu Lys Tyr Phe Gln Gly Ile Lys Glu Val Ile Arg
115 120 125
Lys Glu Phe Lys Phe Lys Gly Glu Pro Asn Pro Lys Asn Ile Glu Met
130 135 140
Val Lys Leu Met Arg Gly Ser Asn Ser Val Ser Ile His Phe Arg Arg
145 150 155 160
Gly Asp Tyr Ile Ser Asn Pro Asp Ala Tyr Lys Val His Gly Gly Ile
165 170 175
Thr Thr Ile His Tyr Tyr Glu Asn Ala Val Lys Glu Ile Lys Ser Lys
180 185 190
Val Lys Glu Pro Lys Phe Phe Ile Phe Ser Asp Asp Ile Lys Trp Val
195 200 205
Lys Glu Asn Phe Lys Leu Glu Asp Ala Phe Phe Ile Asp Trp Asn Thr
210 215 220
Gly Ser Glu Ser Tyr Arg Asp Ile Glu Leu Met Ser Asn Cys Lys His
225 230 235 240
Asn Ile Ile Ala Asn Ser Thr Phe Ser Trp Trp Gly Ala Trp Leu Asn
245 250 255
Lys Asn Lys Asn Lys Ile Val Ile Ala Pro Asn Gln Trp Phe Asn Thr
260 265 270
Ile Asp Thr Glu Asp Val Ile Pro Asp Thr Trp Gln Cys Ile Asn Thr
275 280 285
Phe
<210> 7
<211> 308
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Bt2FT
<400> 7
Met Asn Glu Ile His Val Leu Leu Thr Gly Arg Leu Gly Asn Gln Leu
1 5 10 15
Phe Gln Tyr Ala Phe Ala Arg Ser Leu Gln Lys Gln Tyr Gly Gly Arg
20 25 30
Ile Val Cys Asn Thr Phe Asp Leu Asp His Arg Ser Glu Lys Ser Lys
35 40 45
Val Ala Asp Gln Lys Phe Ser Tyr Ala Met Gly Asp Phe Lys Leu Asp
50 55 60
Glu Asn Val Ala Leu Glu Asp Ala Ala Leu Pro Trp Tyr Ala Asp Phe
65 70 75 80
Ser Ser Pro Leu Ile Lys Pro Val Lys Lys Leu Phe Pro Arg Arg Tyr
85 90 95
Phe Asp Phe Met Ala Arg Arg Gly Tyr Leu Met Trp Gln Arg Thr Asp
100 105 110
Tyr Met Pro Ile Pro Lys Leu Glu Cys Glu Asp Val Phe Ala Trp Gly
115 120 125
Trp Trp Gln Asp Ile Arg Tyr Phe Gln Asp Val Gln Thr Glu Leu Ser
130 135 140
Asp Glu Val Val Pro Val Thr Asp Pro Leu Pro Glu Asn Gln Tyr Ile
145 150 155 160
Tyr Asn Ala Ala Ser Gly Glu Glu Ser Val Cys Ile Ser Ile Arg Cys
165 170 175
Gly Asn Tyr Phe Asn Pro Thr Val Lys Lys Leu Leu Tyr Val Cys Thr
180 185 190
Pro Glu Tyr Phe Arg Asn Ala Val Glu Ser Ile Thr Lys Arg Leu Thr
195 200 205
His Pro Lys Phe Ile Val Phe Thr Asp Asp Val Asp Trp Val Lys Glu
210 215 220
Asn Ile Arg Phe Glu Ser Ala Tyr Pro Gln Tyr Glu Phe Leu Tyr Glu
225 230 235 240
Arg Gly Ser Asp Thr Val Glu Glu Lys Ile Arg Met Met Thr Leu Cys
245 250 255
Lys His Phe Ile Ile Ser Asn Ser Thr Phe Ser Trp Trp Ala Gln Phe
260 265 270
Leu Ser Lys Ser Glu Asn Lys Ile Val Ile Ala Pro Asp Arg Trp Phe
275 280 285
Val Asp Gly Arg Arg Ile Gly Leu Tyr Met His Gly Trp Thr Leu Ile
290 295 300
Pro Ala Gly Gln
305
<210> 8
<211> 317
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Fp2FT
<400> 8
Met Ile Tyr Val Glu Met His Gly Arg Leu Gly Asn Gln Met Phe Gln
1 5 10 15
Tyr Ala Ala Ala Arg Ala Leu Gln Glu Lys Asn Asn Gln Pro Ile Met
20 25 30
Leu Ser Phe Arg Lys Val Ile Gly Ala Asn Thr Glu Gly Thr Ala Gly
35 40 45
Trp Glu Asn Ser Leu Lys Tyr Phe Asn Val Lys Pro Cys Glu Tyr Tyr
50 55 60
Met Gly Lys Lys Ser Leu Val Thr Glu Tyr Pro Val Glu Tyr Arg Leu
65 70 75 80
Leu Cys Tyr Ala Tyr Ala Leu Ser Tyr Lys Pro Leu Met Asn Asn Met
85 90 95
Asn Arg Trp Tyr Glu Tyr Gln Val Lys Cys Cys Arg Phe Leu Asp Arg
100 105 110
Phe Gly Ile Arg Trp Ile Ala Asn Gly Tyr Tyr Asp Phe His Tyr Asn
115 120 125
Gly Leu Lys Asn Tyr Leu Leu Asn Gly Ser Phe Glu Ser Pro Lys Tyr
130 135 140
Phe Asp Ser Ile Arg Asp Lys Leu Leu Glu Glu Phe Thr Pro Arg Glu
145 150 155 160
Glu Glu Arg Lys Glu Asn Lys Arg Leu Tyr Glu Gln Ile Arg Lys Arg
165 170 175
Asn Ser Val Cys Leu Ser Val Arg His Phe Gln Leu Thr Gly Lys Gln
180 185 190
Ala Asp Met Tyr Asp Val Cys Ser Leu Glu Tyr Tyr Gln Thr Ala Ile
195 200 205
Arg Lys Met Cys Glu Leu Ile Glu Asn Pro Leu Phe Val Val Phe Ser
210 215 220
Asp Asp Ile Glu Trp Val Lys Asn Thr Ile Asp Leu Ser Arg Val Glu
225 230 235 240
Val Val Tyr Glu Thr Pro Gly Asn Pro Val Trp Glu Lys Leu Arg Leu
245 250 255
Met Tyr Ser Cys Lys Asn Phe Ile Ile Pro Asn Ser Thr Phe Ala Trp
260 265 270
Trp Ala Gln Tyr Leu Ser Arg Asn Pro Asp Lys Tyr Val Leu Cys Pro
275 280 285
Ala Lys Trp Phe Asn Asn Asn Phe Glu Ser Pro Leu Ile Ala Ser Gln
290 295 300
Trp Val Arg Ile Asp Arg Glu Gly Asn Ile Val Asn Glu
305 310 315
<210> 9
<211> 289
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Gs2FT
<400> 9
Met Lys Ile Val Lys Val Ile Gly Gly Leu Gly Asn Gln Met Phe Gln
1 5 10 15
Tyr Ala Phe Tyr Arg Asn Leu Lys Ala Lys Phe Gln Glu Val Lys Leu
20 25 30
Asp Ile Thr Ala Phe Glu Thr Tyr Lys Leu His Asn Gly Tyr Glu Leu
35 40 45
Glu Arg Val Phe Asp Ile Lys Pro Glu Tyr Ala Thr Lys Lys Glu Ile
50 55 60
Tyr Pro Leu Thr Thr Asn Arg Asn Ser Lys Ile Ser Lys Ile Lys Arg
65 70 75 80
Arg Ile Phe Gly Gly Lys Glu Thr Glu Tyr Ile Glu Lys Asp Leu Lys
85 90 95
Phe Asp Pro Glu Val Phe Lys Val Thr Gly Asp Val Tyr Phe Glu Gly
100 105 110
Tyr Trp Gln Thr Glu Lys Tyr Phe Lys Glu Ile Glu Asp Leu Ile Arg
115 120 125
Lys Asp Phe Gln Phe Lys Asn Pro Leu Thr Asn Lys Asn Leu Glu Leu
130 135 140
Ser Asn Lys Ile Lys Asn Glu Asn Ser Val Ser Ile His Val Arg Arg
145 150 155 160
Gly Asp Tyr Tyr Thr Ser Lys Lys Ala Glu Arg Lys His Gly Asn Ile
165 170 175
Ala Thr Ile Glu Tyr Tyr Gln Lys Ala Val Arg Lys Ile Thr Glu Phe
180 185 190
Val Asp Asn Pro Val Phe Tyr Ile Phe Ser Asp Asp Ile Pro Trp Val
195 200 205
Lys Glu Asn Leu Lys Leu Glu Asn Glu Val Ile Tyr Val Asp Trp Asn
210 215 220
Lys Gly Leu Asp Ser Tyr Ile Asp Met Gln Leu Met Ser Ile Cys Lys
225 230 235 240
His Asn Ile Ile Ala Asn Ser Thr Phe Ser Trp Trp Gly Ala Trp Leu
245 250 255
Asn Gln Asn Lys Asn Lys Ile Val Ile Ala Pro Ser Arg Trp Ile Asn
260 265 270
Asn Lys Arg Leu Asp Thr Ser Asp Val Ile Pro Lys Glu Trp Ile Lys
275 280 285
Ile
<210> 10
<211> 314
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Lg2FT
<400> 10
Met Leu Tyr Val Glu Met Asp Gly Arg Cys Gly Asn Gln Leu Phe His
1 5 10 15
Tyr Ala Val Ala Arg Tyr Ile Gln Leu Ala Ile Gly Asn Lys Glu Lys
20 25 30
Leu Cys Leu Asn Phe Asn Lys Ile Phe Glu Lys Lys Asp Glu Asn Asn
35 40 45
Gly Trp Val Asp Tyr Leu Lys Asp Phe Lys Thr Val Pro Tyr Ser Tyr
50 55 60
Tyr Ser Lys Ser Gly Thr Ile Leu Lys Asn Glu Ser Asn Phe Ile Gln
65 70 75 80
Lys Ile Ala Ile Gly Leu Lys Ala Ile Gln Ile Lys Ser Leu Thr Lys
85 90 95
Lys Ser Arg Gln Glu Gln Ala Asp Lys Ala Glu Val Gly Gln Arg Thr
100 105 110
Leu Asn Lys Leu Gly Val Tyr Trp Val Arg Glu Gly Val Asn Gln Ile
115 120 125
Tyr Pro Tyr Lys Asn Asn Lys Ile Leu Val Ser Gly Ile Cys Glu Ser
130 135 140
Asn Phe Ile Tyr Glu Ile Gln Glu Gln Leu Gln Lys Glu Leu Ile Pro
145 150 155 160
Val Thr Pro Val Ser Ser Leu Asn Lys Ser Leu Leu Glu Lys Ile Asp
165 170 175
Asn Cys Asn Ser Val Cys Ile Ser Val Arg Arg Gly Asp Phe Phe Asn
180 185 190
Asn Lys Asn Ala Lys Lys Tyr Gly Val Cys Ser Pro Glu Tyr Tyr Ile
195 200 205
Arg Ala Lys Lys Tyr Phe Asp Lys Lys Arg Leu Glu Asn Thr Val Tyr
210 215 220
Phe Cys Phe Ser Asp Asp Ile Glu Trp Cys Lys Glu Asn Leu Lys Phe
225 230 235 240
Thr Asp Lys Asn Val Ile Phe Val Ser Gln Glu Met Pro Val Tyr Glu
245 250 255
Thr Leu Arg Leu Met Ser His Cys Lys His Phe Ile Leu Ser Asn Ser
260 265 270
Thr Phe Ser Trp Trp Gly Gln Phe Leu Ser Glu Tyr Lys Asp Lys Ile
275 280 285
Val Val Ser Pro Ala Arg Trp Asn Asn Asp Gly Tyr Asp Thr Asn Leu
290 295 300
Ile Asp Lys Asn Trp Ile Leu Ile Asp Ala
305 310
<210> 11
<211> 309
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Ll2FT
<400> 11
Met Ile Tyr Val Glu Ile Arg Gly Asn Leu Gly Asn Gln Leu Phe Ile
1 5 10 15
Tyr Ala Thr Ala Lys Lys Ile Gln Lys Leu Thr Gly Gln Lys Ile Gln
20 25 30
Leu Asn Thr Thr Thr Leu Asn Lys Tyr Phe Pro Asn Tyr Lys Phe Gly
35 40 45
Leu Ser Glu Phe Ile Met Glu Asp Pro Asp Cys Phe Ile Glu Ser Tyr
50 55 60
Lys Lys Leu Pro Trp Phe Thr Asn Glu Tyr Leu Leu Pro Ile Lys Ile
65 70 75 80
Phe Lys Lys Ile Leu Asn Lys Thr Pro Lys Ile Asn Lys Ile Leu Ser
85 90 95
Asp Phe Phe Phe Lys Ala Phe Glu Lys Lys Gly Tyr Phe Ile Trp Arg
100 105 110
Gly Glu Thr Phe Lys Lys Phe Ser Leu Gly Asn His Lys Asn Tyr Tyr
115 120 125
Leu Ser Gly Phe Trp Gln Ser Glu Asp Tyr Phe Tyr Asp Ile Arg Asp
130 135 140
Glu Leu Leu Glu Ile Ile Thr Pro Ile Asn Ser Ile Arg Glu Cys Asn
145 150 155 160
Phe Glu Leu Leu Asn Leu Ile Arg Asn Ser Glu Ser Ile Cys Val Ser
165 170 175
Ile Arg Arg Gly Asp Tyr Val Asp Asn Pro Lys Ile Ser Ala Ile Tyr
180 185 190
Asn Val Cys Asp Ile Asn Tyr Phe Ile Glu Ser Val Asn Glu Ile Lys
195 200 205
Lys Asn Val Val Asn Val Lys Val Ile Cys Phe Ser Asp Asp Val Glu
210 215 220
Trp Val Lys Lys Asn Ile Lys Phe Asp Cys Glu Thr His Tyr Glu Thr
225 230 235 240
Tyr Gly Asn Ser Leu Ser Glu Lys Val Gln Leu Met Ser Ser Cys Lys
245 250 255
His Phe Val Leu Ser Asn Ser Ser Phe Ser Trp Trp Thr Glu Phe Leu
260 265 270
Ser Ile Arg Gly Gly Ile Thr Ile Ala Pro Lys Asn Trp Tyr Ala Asp
275 280 285
Glu Arg Glu Ala Asp Ile Tyr Arg Lys Asn Trp Ile Tyr Leu Glu Asp
290 295 300
Lys Thr Glu Glu Glu
305
<210> 12
<211> 306
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Ls2FT
<400> 12
Met Glu Glu Ile His Thr Leu Leu Thr Gly Arg Leu Gly Asn Gln Leu
1 5 10 15
Phe Gln Tyr Ala Phe Ala Arg Asn Leu Gln Lys Gln Tyr Gly Gly Gln
20 25 30
Ile Tyr Cys Asp Val Tyr Glu Leu Glu His Arg Met Ser Lys Val Ala
35 40 45
Asp Glu Lys Phe Ser Tyr Ala Met Ser Gly Phe Lys Leu Asp Ala Gly
50 55 60
Val Ile Arg Glu Asp Gln Ala Phe Pro Trp Tyr Ala Asp Phe Ser Asn
65 70 75 80
Pro Val Ile Lys Pro Ile Lys Lys Ala Met Pro Arg Lys Tyr Phe Glu
85 90 95
Leu Met Ala Arg Arg Gly Tyr Leu Met Trp Gln Arg Ser Asp Tyr Met
100 105 110
Pro Ile Pro Val Leu Asp Thr Gln Arg Val Phe Ala Ser Gly Trp Trp
115 120 125
Gln Asp Ile Arg Tyr Leu Gln Asn Val Gln Glu Glu Leu Ser Asp Glu
130 135 140
Ile Val Pro Ile Thr Asn Pro Leu Gln Glu Asn Arg Tyr Ile Tyr Asp
145 150 155 160
Ala Ala His Asp Arg Asp Ser Val Cys Ile Ser Ile Arg Cys Gly Asn
165 170 175
Tyr Phe Asn Pro Thr Val Lys Lys Leu Leu Tyr Val Cys Asn Pro Gln
180 185 190
Tyr Phe His Asp Ser Val Lys Arg Ile Ser Gln Met Leu Ala His Pro
195 200 205
Lys Phe Ile Val Phe Thr Asp Asp Val Ser Trp Val Lys Glu His Leu
210 215 220
Lys Phe Glu Asp Thr Tyr Pro Gln Phe Glu Phe Leu Tyr Glu Arg Gly
225 230 235 240
Cys Asp Thr Ala Glu Glu Lys Ile Arg Met Met Ala Met Cys Asn Asn
245 250 255
Phe Ile Ile Ser Asn Ser Thr Phe Ser Trp Trp Ala Gln Phe Leu Ser
260 265 270
Lys Asn Lys Glu Lys Ile Val Ile Ala Pro Asp Lys Trp Phe Val Asp
275 280 285
Gly Arg Lys Ile Gly Leu Tyr Met Asp Gly Trp Thr Leu Val Pro Ala
290 295 300
Gly Arg
305
<210> 13
<211> 292
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of Ri2FT
<400> 13
Met Ile Ala Val Lys Ile Gly Asp Gly Met Gly Asn Gln Leu Phe Asn
1 5 10 15
Tyr Ala Cys Gly Tyr Ala Gln Ala Arg Arg Asp Gly Asp Ser Leu Val
20 25 30
Leu Asp Ile Ser Glu Cys Asp Asn Ser Thr Leu Arg Asp Phe Glu Leu
35 40 45
Asp Lys Phe His Leu Lys Tyr Asp Lys Lys Glu Ser Phe Pro Asn Arg
50 55 60
Asn Leu Gly Gln Lys Ile Tyr Lys Asn Leu Arg Arg Ala Leu Lys Tyr
65 70 75 80
His Val Ile Lys Glu Arg Glu Val Tyr His Asn Arg Asp His Arg Tyr
85 90 95
Asp Val Asn Asp Ile Asp Pro Arg Val Tyr Lys Lys Lys Gly Leu Arg
100 105 110
Asn Lys Tyr Leu Tyr Gly Tyr Trp Gln His Leu Ala Tyr Phe Glu Asp
115 120 125
Tyr Leu Asn Glu Ile Thr Ala Met Met Thr Pro Ala Tyr Glu Gln Ser
130 135 140
Glu Thr Val Lys Lys Leu Gln Glu Glu Phe Lys Lys Thr Pro Thr Cys
145 150 155 160
Ala Val His Val Arg Gly Gly Asp Ile Met Gly Pro Ala Gly Ala Tyr
165 170 175
Phe Lys His Ala Met Glu Arg Met Glu Gln Glu Lys Pro Gly Val Arg
180 185 190
Tyr Ile Val Phe Thr Asn Asp Met Glu Arg Ala Glu Glu Ala Leu Ala
195 200 205
Pro Val Leu Glu Ser Gln Lys Lys Asp Ala Val Gly Gln Ala Glu Asn
210 215 220
Arg Leu Glu Phe Val Ser Glu Met Gly Glu Phe Ser Asp Val Asp Glu
225 230 235 240
Phe Phe Leu Met Ala Ala Cys Gln Asn Gln Ile Leu Ser Asn Ser Thr
245 250 255
Phe Ser Thr Trp Ala Ala Tyr Leu Asn Gln Asn Pro Asp Lys Thr Val
260 265 270
Ile Met Pro Asp Asp Leu Leu Ser Glu Arg Met Arg Gln Lys Asn Trp
275 280 285
Ile Ile Leu Lys
290
<210> 14
<211> 786
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding Am2FT_2
<400> 14
atgaacatgg aacggaaaac cggccttatg aataaaaaat atgtttcccc gtgtttcctg 60
cccggcatgc ggctgggcaa catcatgttc acgctggcgg ccgcctgtgc ccacgcccgt 120
accgtagggg tggagtgccg ggtgccctgg gcttacaatg acgcttccct gatgctgcgc 180
tcccggctcg gcggctgggt gctcccttct acgccgtgcg gcaccaatga acctccatcc 240
tggcaggaac cttcctttgc ctattgtccg gttccctccc gcatccggac gggcgggctt 300
cgcggctact tccagagcgc acggtatttt gaagggcagg aagcgttcat ccgggcgctt 360
ttcgcgcctc tcaccgcgga gaaggagccc ggcgccgttg gcattcacat ccggcttggg 420
gactacaggc gcctccgtga caagcaccgc attctggacc ctggtttcct gcggagggcc 480
gcagggcatt tatcatccgg gaagaaccgt ctggtgctgt tcagcgatga gccggacgag 540
gcggcggaga tgcttgcgcg cgtgccggca ttcggacgct ttgcgctgga aatcgaccgc 600
ggcgctccgt gcgaatccct ccgccgcatg acggcgatgg aggaactcgt catgtcgtgt 660
tcttcctttt cctggtgggg cgcgtggctg ggaaataccc ggaaggtcat cgttccgcgc 720
gactggtttg tcggcggtgt ggaagactac cgggatattt acctgcccca ttgggtaaca 780
ctgtaa 786
<210> 15
<211> 876
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding Bm2FT
<400> 15
atgaaaattg ttcagatttc ttctggtctg ggtaaccagc tgttccagta cgccctgtac 60
aaacgtctgt ctatgaacaa caacgatgtg ttcctggatg tggagacgtc ttatcaactg 120
aacaaaaatc agcacaatgg ttacgaaatt gaacgtattt tttccatcca gccatcccat 180
gctaccaagg gtatgatcga tgaactggcc gatgtagaca accgtctgat caaccgctta 240
cgtcgtaaac tgtttggtcc gaaaaactct atgtataccg aaaccaaaga gttcagctac 300
gattgtgaag ttttcaccaa agatggtatc tatatcaagg gctactggca gaactataac 360
tattttaaag agattgaaga cgatctgaaa aatgaactgg tctttaaaaa agctctggat 420
ctgaaaaatt cgcacctgat caatcagatg aataaagaaa ttagcgtaag catccacgta 480
cgccgcggcg attattacct gaacaaggag tacgaaaaca aatttggcaa cattgcggat 540
cttgattatt atctgaaagc gattaacttc atcaaaaaag aagtagacga tcccaaattt 600
tacgttttct ccgacgatat taaatgggct aaagaaaatc tgaacctgac ggataacgta 660
acttatgttg aacacaacaa agggagcgac tcttacaaag acatgcgtct gatgacctgc 720
tgtaaacata acatcatcgc caactctaca ttctcttggt ggggcgcgtt tcttaacgag 780
aacaagaaca aaatcgtaat cgcgccgggc aaatggatca acgttgaagg cgtcggtggc 840
atcaacctgt tccctgaagg ttggatcgtt tactga 876
<210> 16
<211> 900
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding Bm2FT_2
<400> 16
atgggttgca taaaaagact attcttatat gaatacggag gacgatgttt tttggtaatt 60
gttaaaataa aaggtggatt tggtaaccaa ttgtttacat acgctagtgc ttatgcaata 120
tctcaagaac tgcaacaaaa cttaattatg gataaggtaa tatatgatct agattacttt 180
agaaaatttg aattaccttc attatcgttg aagtatgatc aaatgttaat tagtaagttt 240
gtacctaata ctaaggttaa aactttaatt tataaagttt taaggaggtt aaaattaaaa 300
ggatttacag aggtacatga aaaaaaggaa ttttcttttg atgaaaatat ttacaatttg 360
tcaggtgata tttatttaga tggttattgg cagaattata gatatttcca taaatattat 420
aaagatctgt cggaaatgtt tgtaccaaga gaaactccta gaaaagaagt aactgattat 480
atcacgtctt tacggggagt taattctgtt gctatgcatg ttaggagagg agattataaa 540
acttttaacg gaggaaaatg tttgtcttta gattattata ttaaggctat ggaatatttc 600
aattcaaatg atgtacaatt ttatgtattt acagatgata ttgatttttg tgagaaaaac 660
ctcccgaata gtgaaaatat taattatgtg tcaagaagcg agaaacttac agatattgaa 720
gaattcttta ttatgaaaga atgtaaaaat tttataattg ctaatagtag tttttcgtgg 780
tgggctgctt atttaagtga acagaaagca gatagtttaa ttgtagcccc tgttgttgac 840
atgtggaaaa gagattttta tccagatgag tgggtggcat tgaatacaca tttagaatag 900
900
<210> 17
<211> 864
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding Bm2FT_3
<400> 17
atgataattg ttcaaatcaa gggcggatta ggtaatcaac ttttttcgta tgcttctgct 60
tatggaattg caagagagaa tgaaacagaa ttaattattg atcgatatat atatgataca 120
agctactctt taagaaaata tatgttggat tttttccctg agatcgatga agcactactt 180
ctaaagtata tacctaaaaa aaataaaatt tcacaaatta tgtataaatt gattagaaaa 240
tctaagttga aatataagta caaagctcaa ttatttttag aagaagagga atttaagttt 300
actagaatta gtacccagaa tgaaaatttg tatttaaatg gttattggca aagttatgtg 360
tactttgata aatatcgtaa tgacattatt aaaaaattca cccctttagt aagctttaat 420
caagatggaa atcaacttct taacgaaatt aaaagttata attcagtggc tatccatgtc 480
aggagagggg attatattaa ttttaaaggt ggaaagtgtt tagatagttc ttattatata 540
aaggcaatga agcatttata caatttaaaa ggaaaaaatt tgttttttta tatttttaca 600
gatgatgtgg aatattgtaa aagaattttt aaaaacgtag ctaatgttaa atttataggt 660
gaggaagcaa aattgagtga ttttgaagag tttactttga tgactcattg taaaaacctc 720
ataataggta acagctcatt ttcttggtgg gcagcttact tagcctcttg taaagataaa 780
gctgtaatag ctccagtggt agatatgtgg actgaagatt tctatctacc tgaatggata 840
aaaattaaag cggatttaca atag 864
<210> 18
<211> 876
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding Bs2FT
<400> 18
atgaaaatag tacacataag tagcggttta ggaaatcaaa tgtttcaata tgctctatat 60
aaaaaactgt ccttaattca agataatgtt tttttggata caattaccag ttatcagttg 120
tatcctaatc aacataatgg atatgagtta gagaaggtat ttacaattaa accaaggcat 180
gcaagtaaag aattaacata caatttatca gacttagata attcagtaac atctcgcatt 240
cgtaggaaat taattggaag taaaaagagc atgtatatag aacataaaga atttgaatac 300
gatcctaatt tattttatca agaaaatata tatataaaag gttactggca gaattatgat 360
tattttaaag atatagaaaa tgaattaaag aatgatttta cttttcaaag agctttagat 420
gagaaaaata ataatttagc tattaaaatt aacaatgaga attcaatatc aattcatgta 480
agacgtggtg attattactt aaacagaaag aatcaagaga agtttggaga tatcgctaat 540
ttggaatact attcaaaggc aattagctat attaaagaaa gaatagacaa tccaaagttt 600
tatatctttt cagataatgt tgagtgggta aagcaaaatc ttaattcgtt agaggaagct 660
gtatatatag attataacgt gggaaatgat agttacaagg atatgcaact aatgagtctt 720
tgtaaacata atattattgc taatagttct tttagttggt ggggagcatt tttgaataag 780
aatatggaga aaatagttat tgcacctggg aagtggatta atatgaaagg tgtaaaaaaa 840
gtcaatcttt tcccaaagga ttggataatt tattaa 876
<210> 19
<211> 462
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of GTP-mannose-1-phosphate
guanylyltransferase
<400> 19
Met Lys Leu Val Leu Leu Ser Gly Gly Ser Gly Lys Arg Leu Trp Pro
1 5 10 15
Leu Ser Asn Asp Thr Arg Ser Lys Gln Phe Leu Lys Val Leu Glu Gly
20 25 30
Arg Asp Gly Gln Lys Glu Ser Met Val Gln Arg Val Trp Ser Gln Ile
35 40 45
Glu Ala Val Gly Leu Asn Glu Ser Thr Val Ile Ala Thr Ser Lys Leu
50 55 60
Gln Arg Asp Met Ile Thr Gly Gln Val Gly Thr Asp Ile Pro Leu Ile
65 70 75 80
Ile Glu Pro Glu Arg Arg Asp Thr Phe Pro Ala Ile Ala Leu Ala Ala
85 90 95
Ser Tyr Leu His Ser Glu Glu Gln Ala Ser Glu Asp Glu Ile Val Val
100 105 110
Val Leu Pro Val Asp Pro Tyr Val Glu Asp Gly Phe Phe Glu Arg Val
115 120 125
Lys Asp Leu Glu Ser Val Leu Arg Asp Ser Lys Ala Asp Leu Ala Leu
130 135 140
Met Gly Val Thr Pro Thr Tyr Pro Ser Ser Lys Tyr Gly Tyr Ile Val
145 150 155 160
Pro Glu Gly Lys Val Gln Gly Asp Ser Tyr Phe Thr Val Asp Cys Phe
165 170 175
Thr Glu Lys Pro Ser Glu Glu Ala Ala Val Gly Leu Ile Asn Gln Arg
180 185 190
Ala Leu Trp Asn Cys Gly Val Phe Ala Phe Arg Leu Gly Tyr Leu Leu
195 200 205
Gln Ile Leu Gln Glu Lys Lys Leu Pro Leu Gln Phe Glu Thr Leu His
210 215 220
Lys Glu Tyr Ala Thr Leu Pro Lys Ile Ser Phe Asp Tyr Glu Val Val
225 230 235 240
Glu Lys Ala Ser His Ile Val Ala Leu Pro Tyr Asp Gly Tyr Trp Lys
245 250 255
Asp Leu Gly Thr Trp Asn Thr Leu Thr Glu Glu Met Ala Thr Ser Gln
260 265 270
Ile Gly Lys Gly Val Leu Thr Asp Asp Ser Ala Asn Thr His Leu Val
275 280 285
Asn Glu Leu Asp Ile Pro Val Thr Val Leu Gly Val Lys Asp Val Val
290 295 300
Val Ala Ala Ser Pro Asp Gly Ile Leu Val Ala Asp Lys Ala Ser Ser
305 310 315 320
Pro Lys Ile Lys Glu Ile Ile Gly Asp Trp Asn Tyr Pro Pro Met Tyr
325 330 335
Glu Glu Arg Leu Trp Gly Trp Ser Arg Val Leu Asp Tyr Ala Lys Tyr
340 345 350
Asp Glu Glu Thr Glu Met Val Thr Lys Arg Ile His Ile Ser Lys Gly
355 360 365
Lys Asn Ser Thr Tyr His Tyr His Asn Leu Arg Asp Glu Val Trp Thr
370 375 380
Ile Val Arg Gly Glu Gly Glu Leu Ala Leu Asp Asp Tyr Ile Arg Arg
385 390 395 400
Val Lys Ala Gly Asp Ile Ile His Leu Pro Ala Gly Lys Lys His Gly
405 410 415
Ile His Ala Leu Thr Asp Leu Glu Phe Ile Glu Val Gln Thr Gly Lys
420 425 430
Val Ile Ser Asn Glu Asp Ser Ile Arg Leu Ile Tyr Asp Trp Asn Asp
435 440 445
Val Ile Lys Gln Phe Gln Ser Ser Arg Pro Gln Val Ile Leu
450 455 460
<210> 20
<211> 359
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of phosphomannomurase
<400> 20
Leu Lys Thr Ile Ala Tyr Tyr Ile Ser Asp Tyr Gly Tyr Gly His Ala
1 5 10 15
Thr Arg Ser Thr Ala Ile Ile Arg Gln Leu Leu Lys Gln Asn Glu Glu
20 25 30
Ile Glu Ile Ile Ile Cys His Ser Phe Ala Ile Glu Phe Leu Arg Gln
35 40 45
Ser Phe Arg Tyr Glu Arg Arg Val Arg Phe Arg Glu Ile Leu Thr Asp
50 55 60
Val Gly Tyr Val Leu Lys Glu Asn Ser Leu Glu Pro Asp Ala Asp Leu
65 70 75 80
Leu Asn Arg Gln Val Ala Ser Tyr Val Ser Ser Phe His Thr Lys Leu
85 90 95
Met Tyr Glu Lys His Phe Met Lys Val Ala Asn Val Ser Phe Val Ile
100 105 110
Ser Asp Ile Ser Pro Leu Gly Ile Ala Gly Ala Tyr Leu Leu Lys Leu
115 120 125
Pro Ser Leu Gly Ile Ser Asn Phe Thr Trp Phe Thr Ala Tyr Glu Gly
130 135 140
Leu Ile Gly Glu Leu Ser Leu Glu Phe Leu Glu Lys Arg Tyr Lys Gln
145 150 155 160
Met Thr Tyr Gln Phe Ser Leu Ala Cys Ser Asp Glu Pro His Trp Gly
165 170 175
Ile Lys Glu Asn Gln Ser Phe Gly Phe Tyr Ser Arg Glu Ile Asn Asp
180 185 190
Ser Glu Val Gln Arg Ile Arg Lys Glu Ile Asp Pro Glu Gly Glu Ser
195 200 205
Tyr Ile Ile Tyr Phe Gly Leu Gly Met Lys Val Asp Ile Gly Thr Leu
210 215 220
Ser Asp Leu Pro Ile Trp Gln Ser Pro Asn Cys Lys Phe Ile Val Ser
225 230 235 240
Ser Asn Val Asp Val Thr His Pro Asn Val Tyr Lys Ile Pro Gln Asp
245 250 255
Glu Thr Glu Thr Gln His Tyr Val Ala Ala Ala Asp Leu Gly Leu Thr
260 265 270
Lys Ala Gly Trp Gly Thr Ile Ser Glu Ala Val Cys Ala Gly Val Pro
275 280 285
Leu Leu Ile Thr Asn Arg Ser Ser Met Lys Glu Asp Arg His Thr Ile
290 295 300
Asp Tyr Leu Val Arg His Gln Leu Cys Asp Leu Ile Glu Trp Lys Glu
305 310 315 320
Leu Glu Gly Leu Val Val Thr Pro Ser Leu Ile Glu Thr Cys Arg Arg
325 330 335
Gln Gln Ile Asn Arg Tyr Lys Asn Glu Ala Met His Ile Ala Glu Glu
340 345 350
Ile Asn Lys Lys Ile Asn Ser
355
<210> 21
<211> 313
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of GDP-L-fucose synthase
<400> 21
Met Asn Lys Asn Ser Lys Ile Tyr Val Ala Gly His Arg Gly Leu Val
1 5 10 15
Gly Ser Ala Ile Val Arg Lys Leu Gln Glu Lys Gly Tyr Ser Asn Leu
20 25 30
Val Tyr Arg Thr Ser Ala Glu Leu Asp Leu Arg Asp Lys Pro Thr Val
35 40 45
Asp Ala Phe Phe Ala Glu Glu Lys Pro Glu Phe Val Phe Leu Ala Ala
50 55 60
Ala Lys Val Gly Gly Ile Val Ala Asn Asn Glu Tyr Pro Ala Glu Phe
65 70 75 80
Ile His Asp Asn Leu Leu Ile Gln Ser Asn Ile Ile Asp Ala Ala Tyr
85 90 95
Arg Asn Asp Val Glu Lys Leu Leu Phe Leu Gly Ser Thr Cys Ile Tyr
100 105 110
Pro Lys Leu Ala Pro Gln Pro Leu Lys Glu Glu Tyr Leu Leu Thr Gly
115 120 125
Glu Leu Glu Pro Thr Asn Glu Pro Tyr Ala Ile Ala Lys Ile Ala Gly
130 135 140
Ile Lys Met Cys Glu Ser Tyr Asn Arg Gln Tyr Gly Thr Lys Tyr Ile
145 150 155 160
Ser Ala Met Pro Thr Asn Leu Tyr Gly Pro Asn Asp Asn Phe Asp Leu
165 170 175
His Thr Ser His Val Leu Pro Ala Leu Leu Arg Lys Phe His Glu Ala
180 185 190
Lys Glu Ser Asn Ala Pro Phe Val Glu Met Trp Gly Thr Gly Thr Pro
195 200 205
Lys Arg Glu Phe Leu Tyr Ser Asp Asp Leu Ala Asp Ala Cys Val Phe
210 215 220
Leu Met Asn Thr Tyr Glu Gly Asn Glu Ile Val Asn Ile Gly Val Gly
225 230 235 240
Glu Asp Ile Thr Ile Lys Glu Leu Gly Glu Lys Val Lys Ala Val Val
245 250 255
Gly Tyr Glu Gly Asn Ile Gln Phe Asp Thr Ser Lys Pro Asp Gly Thr
260 265 270
Pro Arg Lys Leu Val Asp Val Thr Lys Leu Asn Ser Leu Gly Trp Lys
275 280 285
Ala Glu Thr Ala Leu Glu Asp Gly Leu Gln Lys Ala Tyr Gln Trp Phe
290 295 300
Leu Asp Asn Glu Leu Ala Ala Ala Lys
305 310
<210> 22
<211> 359
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of GDP-D-mannose-4,6-dehydratase
<400> 22
Met Lys Thr Ala Leu Ile Thr Gly Val Thr Gly Gln Asp Gly Ser Tyr
1 5 10 15
Leu Ala Glu Leu Leu Leu Glu Lys Gly Tyr Glu Val His Gly Leu Ile
20 25 30
Arg Arg Ser Ser Ser Tyr Asn Gln Glu Arg Leu Glu Asp Leu Leu Thr
35 40 45
Glu Glu Ala Ala Ser Ala Leu Leu Asn Asn Ser Asn Phe His Leu His
50 55 60
Tyr Gly Asp Val Thr Asp Ala Leu Asn Val Thr Arg Ile Ile Gly Asp
65 70 75 80
Ile Gln Pro Asp Glu Ile Tyr Asn Leu Ala Ala Gln Ser His Val Arg
85 90 95
Val Ser Phe Asp Met Pro Gly Tyr Thr Leu Asp Val Asp Ala Lys Gly
100 105 110
Thr Leu Asn Ile Leu Glu Ala Val Arg Ile Leu Gly Leu Thr Asp Lys
115 120 125
Thr Arg Val Tyr Gln Ala Ser Thr Ser Glu Leu Tyr Gly Lys Val Gln
130 135 140
Glu Val Pro Gln Lys Glu Thr Thr Pro Phe Tyr Pro Arg Ser Pro Tyr
145 150 155 160
Gly Val Ala Lys Ile Tyr Gly Phe Trp Ile Thr Lys Asn Tyr Arg Glu
165 170 175
Ser Tyr Ser Met Phe Ala Val Asn Gly Ile Leu Phe Asn His Glu Ser
180 185 190
Glu Arg Arg Gly Glu Thr Phe Val Thr Arg Lys Ile Thr Leu Ala Ala
195 200 205
Ser Arg Ile Ala Gln Gly Lys Gln Gln Lys Leu Thr Leu Gly Asn Leu
210 215 220
Asp Ser Leu Arg Asp Trp Gly Tyr Ala Lys Asp Tyr Val Glu Cys Met
225 230 235 240
Trp Leu Ile Leu Gln His Asp Lys Pro Glu Asp Phe Val Ile Ala Thr
245 250 255
Gly Glu Met His Ser Val Arg Glu Phe Val Gln Leu Ala Phe Lys His
260 265 270
Thr Gly Ile Glu Ile Glu Trp Ser Gly Glu Gly Leu Glu Glu Lys Gly
275 280 285
Thr Asn Lys Ala Thr Gly Glu Val Ile Val Glu Ile Asp Pro Lys Phe
290 295 300
Phe Arg Pro Ala Glu Val Asp Gln Leu Leu Gly Asp Pro Thr Lys Ala
305 310 315 320
Lys Thr Leu Leu Lys Trp Asn Pro Thr Lys Thr Ser Phe Glu Glu Leu
325 330 335
Val Arg Ile Met Val Glu Ala Asp Met Lys Lys Val Glu Ser Glu Glu
340 345 350
Lys Ile Lys Val Met Phe Asp
355
<210> 23
<211> 1389
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding GTP-mannose-1-phosphate
guanylyltransferase
<400> 23
atgaaacttg tattgttatc gggtggatct ggtaagcgtc tatggccgtt atcaaatgat 60
acgcgttcga agcagttttt aaaggtatta gaaggacgag atggacagaa agagtcaatg 120
gttcagcgcg tctggagtca aattgaagcg gttgggttaa atgagtcaac ggttattgcg 180
acgtctaagc ttcagcgtga tatgattaca ggtcaggttg gcaccgatat tccgttgatt 240
attgaacctg agcgccgcga tacgtttcca gcgattgctt tagctgcttc atatcttcat 300
tccgaagagc aagcatctga agatgaaatc gttgtggtgt tgccagttga cccttatgtg 360
gaagacggat tttttgaacg ggtaaaagat ttagaaagtg tgcttcgtga ttcaaaagcg 420
gatttagcgt taatgggtgt tacacctacg tatccttcat ctaagtacgg gtacatcgtg 480
ccggaaggaa aggtacaagg ggacagttat tttacagtgg attgttttac agaaaagccg 540
agtgaagaag cagctgttgg attaattaat cagcgtgcgc tttggaactg cggcgttttt 600
gcgtttcgtt taggttattt gcttcaaatt ttacaagaga aaaaacttcc acttcaattt 660
gaaacgctac ataaagaata tgcaacattg cctaaaatta gttttgacta tgaagtggtg 720
gaaaaggcaa gtcatattgt cgcgcttccg tatgatgggt actggaaaga tcttggtaca 780
tggaatacgt taacggaaga aatggcgaca tctcaaattg gaaaaggtgt attaacggac 840
gactcagcta atacgcattt agtgaatgaa ttagatattc cggttacggt attaggcgta 900
aaagatgtag ttgttgccgc tagtccagat gggatcttag ttgcggataa agcatctagt 960
cctaagatta aagaaattat tggcgactgg aactaccctc caatgtatga agaacgtctt 1020
tggggctggt cacgcgtgtt agattatgca aagtatgatg aagaaactga aatggtgaca 1080
aaacgtattc atatttcaaa aggaaagaac tcgacgtatc actatcataa tttacgtgat 1140
gaagtatgga cgattgtacg tggagaagga gagcttgcgc ttgatgacta cattcgccgc 1200
gtaaaagcag gggatattat tcacttgccg gctggcaaaa agcacggaat tcatgcgtta 1260
accgaccttg aatttattga agtacaaacg ggaaaagtga tttcaaatga agatagtatt 1320
cgccttattt atgactggaa tgatgtgatc aagcaattcc aatcatcaag accgcaggtg 1380
attttataa 1389
<210> 24
<211> 1080
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding phosphomannomurase
<400> 24
ttgaaaacaa ttgcctatta catttcagac tacggatatg gccacgcgac tagaagcacg 60
gccattattc gtcaattatt aaagcaaaat gaagaaatag aaattattat ttgtcattca 120
tttgcgattg agtttctcag acaatctttt agatatgaac gtcgtgtgag gtttcgagag 180
attttaacgg atgtcggcta cgtgctaaaa gaaaattcac tagaaccaga cgctgatttg 240
ttgaatcgtc aagttgcttc ttatgtatca agctttcata cgaagcttat gtatgagaag 300
cactttatga aagtagccaa cgtttcattc gtgatttcgg atatttctcc gttagggatc 360
gcaggtgcct atctattaaa attgccttct ttaggaatat caaatttcac ttggtttaca 420
gcatacgaag ggttaattgg agagctatcg ctggaatttt tagaaaaacg ttataagcag 480
atgacgtatc agttttcgct agcatgtagt gatgagccac attggggtat caaagaaaat 540
caaagctttg gtttttattc aagagaaata aacgattctg aagtacagcg tattcgtaaa 600
gagattgatc ctgaaggtga gagttacatc atctacttcg gtctgggtat gaaagtagat 660
attggaacgt tgtcggactt gcctatttgg caaagtccaa actgcaagtt tatcgtttca 720
tcgaatgttg atgttactca tcccaatgtt tataaaattc ctcaagatga aacagaaacg 780
cagcactatg tagcagcagc tgatcttggt ttaacaaaag caggctgggg cacgattagt 840
gaagcggttt gcgctggtgt gccgctgttg attacaaatc gcagcagcat gaaagaagat 900
cgtcatacga ttgattattt agttcgtcat cagctatgcg acttaataga gtggaaagag 960
ttagaaggtt tagtagtaac tccttcgctt attgagacat gcaggcggca gcaaatcaat 1020
cgctataaaa atgaagcgat gcatatcgcc gaagaaatca ataaaaaaat aaacagttag 1080
1080
<210> 25
<211> 942
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding GDP-L-fucose synthase
<400> 25
atgaacaaaa attcgaaaat ttacgtagct ggtcatagag gtctcgtcgg ttcagctatc 60
gtacgaaaac tacaagaaaa aggctatagc aatctagtgt atcgcacaag cgctgagcta 120
gatttaagag acaagccgac agtagacgca ttctttgctg aagaaaagcc agaatttgtt 180
ttcttagcag ccgctaaagt aggtggtatt gtggcaaaca atgaatatcc agcggaattt 240
attcacgata acttattgat tcaatcaaat attattgatg cagcatatcg caatgatgta 300
gagaaactat tattcctagg aagtacatgt atttatccaa aattagcgcc tcagccatta 360
aaagaagaat acttactaac aggtgagctg gagccaacaa acgaaccgta tgcaattgct 420
aaaattgctg gaatcaaaat gtgtgaatcc tataaccgtc agtacggaac aaaatatatc 480
tcggcaatgc ctactaacct ttacggtcca aacgacaatt ttgaccttca tacatcacac 540
gttcttcctg cactgcttcg taagttccat gaagcgaaag aaagcaatgc gccgtttgtt 600
gaaatgtggg gaacaggcac accaaaacgt gaatttctat attcagatga cctagcagat 660
gcgtgcgtat tcttaatgaa tacatatgaa ggcaatgaaa tcgttaatat cggtgtggga 720
gaagatatta cgattaaaga acttggcgaa aaagtaaaag cagttgttgg ttatgaagga 780
aatattcagt ttgatacgtc aaaaccagac ggaacgcctc gtaaactagt agacgtaaca 840
aaattgaaca gtcttggctg gaaagcagaa acagctttag aagacggtct tcaaaaagcg 900
tatcagtggt ttttagataa cgaactagct gcagcaaaat aa 942
<210> 26
<211> 1080
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding GDP-D-mannose-4,6-dehydratase
<400> 26
atgaaaacag cattaattac aggtgtaaca ggacaagatg gatcatactt agcagaactt 60
ttactagaaa aaggctatga agtgcacgga ctaattcgcc gcagctcttc ttataaccaa 120
gagcgtctag aagatctttt aacagaagaa gcagcgagcg cattattaaa caactcaaac 180
ttccaccttc actatggtga cgtaacagat gcattaaatg ttactcgtat tattggtgac 240
attcagccag atgaaattta caatctagct gcacagtcac atgtacgtgt gtcttttgat 300
atgcctggat atacattaga tgttgatgca aaaggtacgt taaacattct tgaagctgta 360
cgtattttag ggttaacaga taaaacacgt gtgtaccaag cttctacatc tgaactgtac 420
ggaaaagttc aagaagtacc acaaaaggaa acaacacctt tctaccctcg ttctccatat 480
ggcgtagcga aaatttacgg cttttggatt acaaagaact accgtgaatc ttatagcatg 540
tttgcagtaa acggtatttt atttaaccat gaatcagagc gccgcggcga aacgttcgta 600
acacgtaaaa ttacgcttgc tgcttcacgt atcgcgcaag gcaagcagca aaaattaacg 660
cttggaaact tagactcttt acgtgactgg ggttacgcaa aagactatgt agaatgcatg 720
tggttaattc ttcaacacga taaacctgaa gattttgtta ttgcgactgg tgaaatgcac 780
tcagtacgtg aatttgtaca gcttgcattc aaacatacag gcattgaaat tgagtggagc 840
ggtgaaggtc tagaagaaaa aggaacaaat aaagcaacgg gtgaagtgat cgtagaaatc 900
gatcctaagt tcttccgtcc tgcagaagtg gatcaattgt taggtgatcc aactaaagcc 960
aagactttat taaagtggaa ccccactaag acttcgtttg aagaattagt tcgcattatg 1020
gtggaagcgg acatgaagaa ggttgaaagc gaagagaaga ttaaggtgat gtttgattaa 1080
1080
<210> 27
<211> 587
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of lactose permease Lac12
<400> 27
Met Ala Asp His Ser Ser Ser Ser Ser Ser Leu Gln Lys Lys Pro Ile
1 5 10 15
Asn Thr Ile Glu His Lys Asp Thr Leu Gly Asn Asp Arg Asp His Lys
20 25 30
Glu Ala Leu Asn Ser Asp Asn Asp Asn Thr Ser Gly Leu Lys Ile Asn
35 40 45
Gly Val Pro Ile Glu Asp Ala Arg Glu Glu Val Leu Leu Pro Gly Tyr
50 55 60
Leu Ser Lys Gln Tyr Tyr Lys Leu Tyr Gly Leu Cys Phe Ile Thr Tyr
65 70 75 80
Leu Cys Ala Thr Met Gln Gly Tyr Asp Gly Ala Leu Met Gly Ser Ile
85 90 95
Tyr Thr Glu Asp Ala Tyr Leu Lys Tyr Tyr His Leu Asp Ile Asn Ser
100 105 110
Ser Ser Gly Thr Gly Leu Val Phe Ser Ile Phe Asn Val Gly Gln Ile
115 120 125
Cys Gly Ala Phe Phe Val Pro Leu Met Asp Trp Lys Gly Arg Lys Pro
130 135 140
Ala Ile Leu Ile Gly Cys Leu Gly Val Val Ile Gly Ala Ile Ile Ser
145 150 155 160
Ser Leu Thr Thr Thr Lys Ser Ala Leu Ile Gly Gly Arg Trp Phe Val
165 170 175
Ala Phe Phe Ala Thr Ile Ala Asn Ala Ala Ala Pro Thr Tyr Cys Ala
180 185 190
Glu Val Ala Pro Ala His Leu Arg Gly Lys Val Ala Gly Leu Tyr Asn
195 200 205
Thr Leu Trp Ser Val Gly Ser Ile Val Ala Ala Phe Ser Thr Tyr Gly
210 215 220
Thr Asn Lys Asn Phe Pro Asn Ser Ser Lys Ala Phe Lys Ile Pro Leu
225 230 235 240
Tyr Leu Gln Met Met Phe Pro Gly Leu Val Cys Ile Phe Gly Trp Leu
245 250 255
Ile Pro Glu Ser Pro Arg Trp Leu Val Gly Val Gly Arg Glu Glu Glu
260 265 270
Ala Arg Glu Phe Ile Ile Lys Tyr His Leu Asn Gly Asp Arg Thr His
275 280 285
Pro Leu Leu Asp Met Glu Met Ala Glu Ile Ile Glu Ser Phe His Gly
290 295 300
Thr Asp Leu Ser Asn Pro Leu Glu Met Leu Asp Val Arg Ser Leu Phe
305 310 315 320
Arg Thr Arg Ser Asp Arg Tyr Arg Ala Met Leu Val Ile Leu Met Ala
325 330 335
Trp Phe Gly Gln Phe Ser Gly Asn Asn Val Cys Ser Tyr Tyr Leu Pro
340 345 350
Thr Met Leu Arg Asn Val Gly Met Lys Ser Val Ser Leu Asn Val Leu
355 360 365
Met Asn Gly Val Tyr Ser Ile Val Thr Trp Ile Ser Ser Ile Cys Gly
370 375 380
Ala Phe Phe Ile Asp Lys Ile Gly Arg Arg Glu Gly Phe Leu Gly Ser
385 390 395 400
Ile Ser Gly Ala Ala Leu Ala Leu Thr Gly Leu Ser Ile Cys Thr Ala
405 410 415
Arg Tyr Glu Lys Thr Lys Lys Lys Ser Ala Ser Asn Gly Ala Leu Val
420 425 430
Phe Ile Tyr Leu Phe Gly Gly Ile Phe Ser Phe Ala Phe Thr Pro Met
435 440 445
Gln Ser Met Tyr Ser Thr Glu Val Ser Thr Asn Leu Thr Arg Ser Lys
450 455 460
Ala Gln Leu Leu Asn Phe Val Val Ser Gly Val Ala Gln Phe Val Asn
465 470 475 480
Gln Phe Ala Thr Pro Lys Ala Met Lys Asn Ile Lys Tyr Trp Phe Tyr
485 490 495
Val Phe Tyr Val Phe Phe Asp Ile Phe Glu Phe Ile Val Ile Tyr Phe
500 505 510
Phe Phe Val Glu Thr Lys Gly Arg Ser Leu Glu Glu Leu Glu Val Val
515 520 525
Phe Glu Ala Pro Asn Pro Arg Lys Ala Ser Val Asp Gln Ala Phe Leu
530 535 540
Ala Gln Val Arg Ala Thr Leu Val Gln Arg Asn Asp Val Arg Val Ala
545 550 555 560
Asn Ala Gln Asn Leu Lys Glu Gln Glu Pro Leu Lys Ser Asp Ala Asp
565 570 575
His Val Glu Lys Leu Ser Glu Ala Glu Ser Val
580 585
<210> 28
<211> 413
<212> PRT
<213> Artificial Sequence
<220>
<223> Amino acid sequence of lactose permease LacY
<400> 28
Met Lys Ser Ser Lys Ser Leu Tyr Trp Lys Leu Ser Ala Tyr Phe Phe
1 5 10 15
Phe Phe Phe Phe Thr Trp Ser Ser Ser Tyr Ser Leu Phe Ser Ile Trp
20 25 30
Leu Gly Gln Glu Ile Lys Leu Asn Gly Ser Ala Thr Gly Leu Ile Phe
35 40 45
Ser Val Asn Ala Ile Phe Ala Leu Cys Met Gln Pro Leu Tyr Gly Tyr
50 55 60
Ile Ser Asp Arg Ile Gly Leu Lys Lys His Val Leu Phe Phe Ile Ser
65 70 75 80
Cys Leu Leu Val Phe Val Gly Pro Phe Tyr Ile Phe Val Tyr Gly Pro
85 90 95
Leu Leu Gln Tyr Asn Val Val Ile Gly Ala Ile Ile Gly Gly Leu Tyr
100 105 110
Leu Gly Val Ala Phe Leu Ala Gly Ile Gly Ala Ile Glu Thr Tyr Ile
115 120 125
Glu Lys Val Ser Arg Lys Tyr Lys Phe Glu Tyr Gly Lys Ser Arg Met
130 135 140
Trp Gly Ser Leu Gly Trp Ala Ala Ala Thr Phe Phe Ala Gly Gln Leu
145 150 155 160
Phe Asn Ile Asn Pro His Ile Asn Phe Trp Val Ala Ser Val Ser Ala
165 170 175
Val Ile Leu Val Ala Ile Ile Phe Ser Val Lys Val Glu Met Ser Ser
180 185 190
Tyr Glu Met Glu Lys Ala Glu Ser Val Thr Leu Arg Asp Val Gly Ser
195 200 205
Leu Phe Leu Leu Lys Glu Phe Trp Phe Phe Met Ile Tyr Val Val Gly
210 215 220
Val Thr Cys Val Tyr Gly Val Tyr Asp Gln Gln Phe Pro Ile Tyr Tyr
225 230 235 240
Ala Ser Leu Phe Pro Thr Glu Ser Ile Gly Asn Gln Val Phe Gly Tyr
245 250 255
Leu Asn Ser Phe Gln Val Phe Leu Glu Ala Gly Met Met Phe Ala Ala
260 265 270
Pro Phe Ile Val Asn Lys Ile Gly Ala Lys Asn Ser Leu Ile Leu Ala
275 280 285
Gly Phe Leu Met Gly Phe Arg Ile Ile Gly Ser Gly Leu Val Val Gly
290 295 300
Pro Ile Gly Ile Ser Ser Met Lys Leu Ile His Ala Leu Glu Leu Pro
305 310 315 320
Ile Met Leu Ile Ala Ile Phe Lys Tyr Leu Ala Ala Asn Phe Asp Thr
325 330 335
Arg Leu Ser Ser Ile Leu Tyr Leu Val Gly Phe Gln Phe Ala Ser Gln
340 345 350
Ile Gly Ala Ser Val Leu Ser Pro Ile Val Gly Gly Leu Tyr Asp Ser
355 360 365
Val Gly Phe Ser Arg Thr Tyr Leu Ile Met Gly Gly Met Val Leu Val
370 375 380
Phe Asn Val Ile Ser Met Phe Thr Leu Leu Asn Ser Lys Lys His Arg
385 390 395 400
Phe Ile Arg Lys Asp Val Gln Glu Asn Thr Gln Ile Ile
405 410
<210> 29
<211> 1764
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding lactose permease Lac12
<400> 29
atggctgatc actcttcttc ttcttcttct ttacaaaaaa aaccaattaa cacaattgaa 60
cacaaagata cattaggtaa cgatcgtgat cacaaagaag ctttaaactc tgataacgat 120
aacacatctg gtttaaaaat taacggtgta ccaattgaag atgctcgtga agaagtatta 180
ttaccaggtt acttatctaa acaatactac aaattatacg gtttatgttt cattacatac 240
ttatgtgcta caatgcaagg ttacgatggt gctttaatgg gttctattta cacagaagat 300
gcttacttaa aatactacca cttagatatt aactcttctt ctggtacagg tttagtattc 360
tctattttca acgtaggtca aatttgtggt gctttcttcg taccattaat ggattggaaa 420
ggtcgtaaac cagctatttt aattggttgt ttaggtgtag taattggtgc tattatttct 480
tctttaacaa caacaaaatc tgctttaatt ggtggtcgtt ggttcgtagc tttcttcgct 540
acaattgcta acgctgctgc tccaacatac tgtgctgaag tagctccagc tcacttacgt 600
ggtaaagtag ctggtttata caacacatta tggtctgtag gttctattgt agctgctttc 660
tctacatacg gtacaaacaa aaacttccca aactcttcta aagctttcaa aattccatta 720
tacttacaaa tgatgttccc aggtttagta tgtattttcg gttggttaat tccagaatct 780
ccacgttggt tagtaggtgt aggtcgtgaa gaagaagctc gtgaattcat tattaaatac 840
cacttaaacg gtgatcgtac acacccatta ttagatatgg aaatggctga aattattgaa 900
tctttccacg gtacagattt atctaaccca ttagaaatgt tagatgtacg ttctttattc 960
cgtacacgtt ctgatcgtta ccgtgctatg ttagtaattt taatggcttg gttcggtcaa 1020
ttctctggta acaacgtatg ttcttactac ttaccaacaa tgttacgtaa cgtaggtatg 1080
aaatctgtat ctttaaacgt attaatgaac ggtgtatact ctattgtaac atggatttct 1140
tctatttgtg gtgctttctt cattgataaa attggtcgtc gtgaaggttt cttaggttct 1200
atttctggtg ctgctttagc tttaacaggt ttatctattt gtacagctcg ttacgaaaaa 1260
acaaaaaaaa aatctgcttc taacggtgct ttagtattca tttacttatt cggtggtatt 1320
ttctctttcg ctttcacacc aatgcaatct atgtactcta cagaagtatc tacaaactta 1380
acacgttcta aagctcaatt attaaacttc gtagtatctg gtgtagctca attcgtaaac 1440
caattcgcta caccaaaagc tatgaaaaac attaaatact ggttctacgt attctacgta 1500
ttcttcgata ttttcgaatt cattgtaatt tacttcttct tcgtagaaac aaaaggtcgt 1560
tctttagaag aattagaagt agtattcgaa gctccaaacc cacgtaaagc ttctgtagat 1620
caagctttct tagctcaagt acgtgctaca ttagtacaac gtaacgatgt acgtgtagct 1680
aacgctcaaa acttaaaaga acaagaacca ttaaaatctg atgctgatca cgtagaaaaa 1740
ttatctgaag ctgaatctgt ataa 1764
<210> 30
<211> 1242
<212> DNA
<213> Artificial Sequence
<220>
<223> Nucleotide sequence encoding lactose permease LacY
<400> 30
atgaaaagtt caaaaagttt atattggaag ctgagtgcat attttttctt cttctttttt 60
acgtggtcgt ccagctactc tttattttct atctggttag ggcaagagat taagttgaac 120
ggatcagcaa caggcttaat cttttctgtg aacgctatct ttgctttgtg tatgcagcct 180
ttgtacggat atatatctga caggataggt ttaaagaaac acgttttatt ttttattagc 240
tgtcttttag tgtttgtagg accattttat atttttgttt atggaccttt gcttcaatac 300
aatgtggtga taggggctat tattggaggc ttatatttag gagtagcgtt tttggcgggc 360
atcggagcga ttgaaacata catagagaaa gtcagccgaa aatataaatt tgagtatgga 420
aaatctagaa tgtgggggtc attaggctgg gctgcagcta cgttttttgc aggacaatta 480
tttaatatta atcctcacat taatttttgg gtggcatctg tttcagcggt tatcctagtc 540
gctattattt tttctgtaaa agtagaaatg agcagttatg aaatggaaaa agctgaatcc 600
gtcacgctaa gagatgtagg tagtttattt ttattgaagg aattttggtt ttttatgatt 660
tatgttgttg gtgttacatg cgtatatggg gtatacgatc agcaatttcc tatctattat 720
gcatctttgt ttcctactga atcgataggt aaccaagtat ttggctactt aaattcgttt 780
caagtatttt tagaagctgg aatgatgttc gcagcaccgt ttattgtaaa taagatcggt 840
gcaaaaaata gtttaatttt agcaggattt ctaatgggat ttcgtattat tgggtcaggg 900
cttgttgttg gtccaatcgg aatatcttca atgaaactca ttcatgcatt agaacttcct 960
attatgctca ttgccatctt taaatatttg gcagctaact ttgacacgcg cctatcttcc 1020
atcctttatt tggttggatt ccaatttgct tcgcaaatag gtgcttcagt tctttcaccc 1080
atagtgggag gattatatga tagcgtaggt ttttcccgta cctatttaat tatgggaggt 1140
atggttttgg tatttaacgt tatttcgatg tttacactac taaactctaa aaaacataga 1200
tttataagaa aagatgtaca agaaaacacg cagattatat ag 1242
<210> 31
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Te2FT_F
<400> 31
catatgatta tcgttcatct gtg 23
<210> 32
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Te2FT_R
<400> 32
ctcgagtcac agaacaatcc acc 23
<210> 33
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bt2FT_F
<400> 33
catatgaacg aaatccacgt gct 23
<210> 34
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bt2FT_1_R
<400> 34
ctcgagtcac tggccagccg gaatc 25
<210> 35
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_F
<400> 35
catatgaaaa ttgttcagat ttct 24
<210> 36
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_R
<400> 36
ctcgagtcag taaacgatcc aacct 25
<210> 37
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Lg2FT_F
<400> 37
catatgctgt acgttgaaat ggacg 25
<210> 38
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Lg2FT_R
<400> 38
ctcgagttac gcgtcgatca ggatc 25
<210> 39
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ls2FT_F
<400> 39
catatggaag agattcacac cctg 24
<210> 40
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ls2FT_R
<400> 40
ctcgagctaa cggccggcag gtac 24
<210> 41
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ll2FT_F
<400> 41
catatgtcta aaaacattta tgtac 25
<210> 42
<211> 26
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ll2FT_R
<400> 42
ctcgagttag attttgatcc agttgt 26
<210> 43
<211> 39
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Am2FT_1_F
<400> 43
aagaaggaga tatacatatg atggctggac attcttgcg 39
<210> 44
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Am2FT_1_R
<400> 44
tggtggtggt ggtgctcgag ttaaatgcgc agccatgagc 40
<210> 45
<211> 39
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Am2FT_2_F
<400> 45
aagaaggaga tatacatatg atgaacatgg aacgtaaga 39
<210> 46
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Am2FT_2_R
<400> 46
tggtggtggt ggtgctcgag ttacagggtc acccagtgcg 40
<210> 47
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Fp2FT_F
<400> 47
aagaaggaga tatacatatg atgatttacg tggaaatg 38
<210> 48
<211> 39
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Fp2FT_R
<400> 48
tggtggtggt ggtgctcgag ttattcgtta acgatgtta 39
<210> 49
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ri2FT_F
<400> 49
aagaaggaga tatacatatg atgatcgcag ttaagatc 38
<210> 50
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Ri2FT_R
<400> 50
tggtggtggt ggtgctcgag ttatttcagg ataatccagt 40
<210> 51
<211> 39
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_2_F
<400> 51
aagaaggaga tatacatatg ggttgcatta aacgcctgt 39
<210> 52
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_2_R
<400> 52
tggtggtggt ggtgctcgag ctattccagg tgagtgttca 40
<210> 53
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_3_F
<400> 53
aagaaggaga tatacatatg atcattgtcc agattaaa 38
<210> 54
<211> 41
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_3_R
<400> 54
tggtggtggt ggtgctcgag ctattgaaga tccgctttga t 41
<210> 55
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_4_F
<400> 55
aagaaggaga tatacatatg atcattgtgc gtgttatc 38
<210> 56
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bm2FT_4_R
<400> 56
tggtggtggt ggtgctcgag ttagaacgta ttaatacact 40
<210> 57
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bs2FT_F
<400> 57
aagaaggaga tatacatatg aaaattgtaa aagttatc 38
<210> 58
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Bs2FT_R
<400> 58
tggtggtggt ggtgctcgag ttagtagatg atccagtctt 40
<210> 59
<211> 36
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Gs2FT_F
<400> 59
aagaaggaga tatacatatg aaaattgtaa aagtta 36
<210> 60
<211> 39
<212> DNA
<213> Artificial Sequence
<220>
<223> primer Gs2FT_R
<400> 60
tggtggtggt ggtgctcgag ttagatttta atccattct 39
<210> 61
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer TAG_PET24ma_F
<400> 61
tagctcgagc accaccacca ccacc 25
<210> 62
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<223> primer TAA_PET24ma_F
<400> 62
taactcgagc accaccacca ccacc 25
<210> 63
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> primer PET24ma_R
<400> 63
catatgtata tctccttctt aaagttaaac aaaattattt 40
Claims (11)
- 서열번호 1 내지 5로 이루어지는 군에서 선택된 1종 이상을 포함하는 α-1,2-푸코오스 전이효소 (α-1,2-fucosyltransferase)가 발현되도록 형질전환된, 재조합 미생물.
- 제1항에 있어서, 상기 재조합 미생물은 GRAS (Generally Recognized As Safe) 미생물인, 재조합 미생물.
- 제1항에 있어서, 상기 재조합 미생물은 바실러스 속 (Bacillus sp.) 미생물, 코리네박테리움 속 (Corynebacterium sp.) 미생물, 대장균 속 (Escherichia sp.) 미생물, 또는 효모 (yeast)인, 재조합 미생물.
- 제3항에 있어서, 상기 바실러스 속 (Bacillus sp.) 미생물은 바실러스 메가테리움 (Bacillus megaterium), 바실러스 서브틸리스 (Bacillus subtilis), 바실러스 세레우스 (Bacillus cereus), 바실러스 코아귤런스 (Bacillus coagulans), 바실러스 리체니포르미스 (Bacillus licheniformis), 또는 바실러스 스테아로테르모필루스 (Bacillus stearothermophilus)이고,
상기 코리네박테리움 속 (Corynebacterium sp.) 미생물은 코리네박테리움 글루타미쿰 (Corynebacterium glutatamicum)이고,
상기 대장균 속 (Escherichia sp.) 미생물은 대장균 (Escherichia coli) 이고,
상기 효모 (yeast)는 사카로미세스 세레비시아 (Saccharomyces cerevisiae), 또는 캔디다 유틸리스 (Candida utilis)인, 재조합 미생물. - 제1항에 있어서, 상기 미생물은 재조합 되기 전 2'-푸코실락토오스 생산능을 가지지 않으며, 재조합에 의해 2'-푸코실락토오스 생산능을 가지는 것인, 재조합 미생물.
- 제1항에 있어서, 상기 재조합 미생물은 푸코오스 합성 유전자를 발현하는 것인, 재조합 미생물.
- 제6항에 있어서, 상기 푸코오스 합성 유전자는 GDP-D-만노오스-4,6-데하이드라타아제 (GDP-D-mannose-4,6-dehydratase), GDP-L-푸코오스 신타아제 (GDP-L-fucose synthase), 포스포만노뮤타아제 (phosphomannomurase), 및 GTP 만노오스-1-포스페이트 구아닐릴트랜스퍼라아제 (GTP-mannose-1-phosphate guanylyltransferase)로 이루어지는 군에서 선택된 1종 이상을 발현하는 것인, 재조합 미생물.
- 제1항에 있어서, 상기 재조합 미생물은 락토오스 막 수송 단백질을 발현하는 것인, 재조합 미생물.
- 제8항에 있어서, 상기 락토오스 막 수송 단백질은 Lac12 및/또는 LacY인, 재조합 미생물.
- 제1항에 있어서, 상기 재조합 미생물은 helicobactor pyori 유래의 α-1,2-푸코오스 전이효소를 포함하는 대조군 대비 2배 이상의 2’-푸코실락토오스 생산성을 가지는 것인, 재조합 미생물.
- 락토오스 (lactose)를 포함하는 배지에서 제1항 내지 제9항 중 어느 한 항에 따른 재조합 미생물을 배양하는 단계를 포함하는, 2’-푸코실락토오스 (2’-fucolsylactose)의 생산방법.
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JP2023540466A JP2024502062A (ja) | 2020-12-31 | 2021-12-28 | フコース転移酵素を発現する組換え微生物およびこれを用いた2’-フコシルラクトース製造方法 |
PCT/KR2021/020001 WO2022145945A1 (ko) | 2020-12-31 | 2021-12-28 | 푸코오스 전이효소를 발현하는 재조합 미생물 및 이를 이용한 2'-푸코실락토오스 제조방법 |
EP21915743.5A EP4273250A1 (en) | 2020-12-31 | 2021-12-28 | Recombinant microorganism expressing fucosyltransferase, and method of producing 2'-fucosyllactose using same |
CN202180093567.6A CN116917485A (zh) | 2020-12-31 | 2021-12-28 | 表达岩藻糖基转移酶的重组微生物和使用其生产2’-岩藻糖基乳糖的方法 |
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CN116200360A (zh) * | 2023-01-30 | 2023-06-02 | 芝诺(苏州)生物科技有限公司 | 一种futCB突变体及生物合成2’-岩藻糖基乳糖的方法 |
WO2024123084A1 (ko) * | 2022-12-06 | 2024-06-13 | (주)에이피테크놀로지 | 베타-갈락토시다아제가 발현되도록 형질전환된 재조합 코리네박테리움 글루타미쿰을 이용한 푸코실락토오스의 생산방법 |
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KR20170035835A (ko) | 2014-05-15 | 2017-03-31 | 글리코신 엘엘씨 | 푸코실레이티드 올리고당의 제조에 사용되는 알파(1,2)푸코실트랜스퍼라아제 동질 유전자형 |
KR101794971B1 (ko) * | 2016-02-02 | 2017-11-08 | 서울대학교 산학협력단 | α-1,3-푸코실락토오즈의 대량 생산 방법 |
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CN116200360A (zh) * | 2023-01-30 | 2023-06-02 | 芝诺(苏州)生物科技有限公司 | 一种futCB突变体及生物合成2’-岩藻糖基乳糖的方法 |
CN116200360B (zh) * | 2023-01-30 | 2023-09-15 | 芝诺(苏州)生物科技有限公司 | 一种futCB突变体及生物合成2’-岩藻糖基乳糖的方法 |
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