KR20160000082A - L-트립토판 생산능을 갖는 에스케리키아속 미생물 및 이를 이용한 l-트립토판의 제조 방법 - Google Patents
L-트립토판 생산능을 갖는 에스케리키아속 미생물 및 이를 이용한 l-트립토판의 제조 방법 Download PDFInfo
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- KR20160000082A KR20160000082A KR1020140076698A KR20140076698A KR20160000082A KR 20160000082 A KR20160000082 A KR 20160000082A KR 1020140076698 A KR1020140076698 A KR 1020140076698A KR 20140076698 A KR20140076698 A KR 20140076698A KR 20160000082 A KR20160000082 A KR 20160000082A
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- leu
- ala
- gly
- val
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- C12P—FERMENTATION OR ENZYME-USING PROCESSES TO SYNTHESISE A DESIRED CHEMICAL COMPOUND OR COMPOSITION OR TO SEPARATE OPTICAL ISOMERS FROM A RACEMIC MIXTURE
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Abstract
본 발명은 L-트립토판 생산능을 갖는 에스케리키아속 미생물에 관한 것으로, 보다 구체적으로 내재적 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 약화시키거나 불활성화시킴으로써, L-트립토판 생산능이 향상된 에스케리키아속 미생물에 관한 것이다.
또한, 본 발명은 상기 에스케리키아속 미생물을 이용하여 L-트립토판을 제조하는 방법에 관한 것이다.
또한, 본 발명은 상기 에스케리키아속 미생물을 이용하여 L-트립토판을 제조하는 방법에 관한 것이다.
Description
본 발명은 L-트립토판 생산능을 갖는 에스케리키아속 미생물, 및 상기 미생물을 이용하여 L-트립토판을 제조하는 방법에 관한 것이다.
L-트립토판(L-tryptophan)은 필수 아미노산의 하나로 사료 첨가제 등으로 널리 사용되어 왔으며, 또한 수액제 등의 의약품 원료 및 건강식품소재 등으로 널리 사용되어 왔다. 이와 같은, L-트립토판은 화학 합성법, 효소 반응법, 발효법 등을 통해 생산될 수 있으나, 현재에는 미생물을 이용한 직접 발효법이 주로 이용되고 있다.
L-트립토판 생산균주의 개발 방향은 초기에는 돌연변이 선별로 진행되거나(대한민국 등록특허 제1987-0001813호), 유전공학의 발달과 함께 생합성 경로 효소들의 트립토판 피드백 저해를 극복하는 방법, 혹은 트립토판 생합성 효소의 발현 강화와 같이 대사과정의 효소 합성 강화를 통해 트립토판 생산 균주의 개발이 진행되었다.
한편, L-아미노산의 생산을 향상시키고자 Entner-Doudoroff 경로를 이용하는 방법이 등장한 바 있다(US 등록특허 제7432085호). 상기 US 등록특허는 Entner-Doudoroff 경로에 관여하는 효소의 활성을 증강시켜 피루브산을 중간물질로 사용하는 생합성 경로에 의해 생산되는 L-아미노산의 생산을 향상시키는 방법에 대한 것으로, 구체적으로 6-포스포글루코네이트 탈수소효소 또는 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 증강시키는 것을 핵심 특징으로 하고 있다.
그러나 본 발명자들은 다른 L-아미노산들과 달리, L-트립토판의 경우에는 상기 Entner-Doudoroff 경로의 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 함께 약화시키거나 불활성화시키는 경우 L-트립토판 생산능이 현저히 향상된다는 것을 최초로 규명하였으며, 이를 이용하여 L-트립토판 생산능이 향상된 에스케리키아속 미생물 및 이를 이용한 L-트립토판 제조 방법에 대한 본 발명을 완성하기에 이르렀다.
본 발명은 L-트립토판 생산능을 갖는 미생물을 제공하는 것을 목적으로 한다.
또한, 본 발명은 상기 L-트립토판 생산능을 갖는 미생물을 이용하여 L-트립토판을 효율적으로 생산하는 방법을 제공하는 것을 목적으로 한다.
상기 과제를 해결하기 위한 일 양태로서, 본 발명은 내재적 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성이 약화 또는 불활성화되어, L-트립토판 생산능을 갖는 에스케리키아속(Escherichia) 미생물을 제공한다.
본 발명에서 사용되는 용어, "L-트립토판(L-tryptophan)"은 α-아미노산의 하나로, 체내에서 합성되지 않는 필수 아미노산이며 화학식이 C11H12N2O2인 방향족 L-아미노산을 말한다.
본 발명에서 사용되는 용어, "Entner-Doudoroff 경로"는 에스케리키아속 미생물 내 존재하는 탄소 대사 경로의 하나로서, 상기 경로로 유입된 탄소원이 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제에 의한 2단계의 일련의 효소 반응을 통하여, 글리세르알데하이드-3-포스페이트(Glyceraldehyde-3-phosphate) 및 피루브산(pyruvate)으로 전환되는 것을 촉매하는 경로이다.
본 발명에서 사용되는 용어, "6-포스포글루코네이트 탈수소효소(6-phosphogluconate dehydratase; Edd; EC 4.2.1.12)"는 Entner-Doudoroff 경로에 관여하는 효소로서, 6-포스포-D-글루코네이트를 2-디하이드로-3-데옥시-6-포스포-D-글루코네이트로 전환시키는 반응을 촉매하는 효소를 의미한다. 상기 효소는 구체적으로 서열번호 1의 아미노산 서열로 나타낼 수 있으나, 상기 효소의 활성을 갖는 서열이라면 제한없이 포함된다. 또한, 상기 6-포스포글루코네이트 탈수소효소를 코딩하는 유전자는 구체적인 예로 서열번호 2의 염기서열로 나타낼 수 있으나, 상기 효소를 코딩할 수 있는 서열이라면 제한없이 포함된다.
본 발명에서 사용되는 용어, "2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제(2-keto-3-deoxy-6-phosphogluconate aldolase; Eda; EC 4.1.2.14)"는 Entner-Doudoroff 경로에 관여하는 효소로서, 2-디하이드로-3-데옥시-6-포스포-D-글루코네이트를 글리세르알데하이드-3-포스페이트 및 피루브산으로 전환시키는 반응을 촉매하는 효소를 의미한다. 상기 효소는 구체적으로 서열번호 3의 아미노산 서열로 나타낼 수 있으나, 상기 효소의 활성을 갖는 서열이라면 제한없이 포함된다. 또한, 상기 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제를 코딩하는 유전자는 구체적인 예로 서열번호 4의 염기서열로 나타낼 수 있으나, 상기 효소를 코딩할 수 있는 서열이라면 제한없이 포함된다.
상기 각 효소들은 상기 서열번호 1 또는 3으로 기재한 아미노산 서열뿐만 아니라, 상기 서열과 70% 이상, 구체적으로는 80% 이상, 더욱 구체적으로는 90% 이상, 보다 더욱 구체적으로는 95% 이상, 더 구체적으로는 98% 이상, 더욱 더 구체적으로는 99% 이상의 상동성을 나타내는 아미노산 서열로서 실질적으로 상기 각 효소와 동일하거나 상응하는 효능을 나타내는 효소라면 제한없이 포함한다. 또한 이러한 상동성을 갖으며 각 효소에 상응하는 효능을 나타내는 아미노산 서열이라면, 일부 서열이 결실, 변형, 치환 또는 부가된 아미노산 서열을 갖는 효소 변이체도 본 발명의 범위 내에 포함됨은 자명하다.
또한, 상기 각 효소들을 코딩하는 유전자는 상기 서열번호 2 또는 4로 기재한 염기서열뿐만 아니라, 상기 서열과 80% 이상, 구체적으로는 90% 이상, 보다 구체적으로는 95% 이상, 더욱 구체적으로는 98% 이상, 보다 더욱 구체적으로는 99% 이상의 상동성을 나타내는 염기 서열로서 실질적으로 상기 각 효소와 동일하거나 상응하는 효능을 나타내는 효소를 코딩하는 유전자 서열이라면 제한없이 포함한다. 또한 이러한 상동성을 갖는 염기서열이라면, 일부 서열이 결실, 변형, 치환 또는 부가된 염기서열도 본 발명의 범위 내에 포함됨은 자명하다.
본 발명에서 용어, "상동성"은 두 개의 폴리뉴클레오티드 또는 폴리펩타이드 모이티 사이의 동일성의 퍼센트를 말한다. 하나의 모이티로부터 다른 하나의 모이티까지의 서열 간 상동성은 알려진 당해 기술에 의해 결정될 수 있다. 예를 들면, 상동성은 서열정보를 정렬하고 용이하게 입수 가능한 컴퓨터 프로그램을 이용하여 두 개의 폴리뉴클레오티드 분자 또는 두 개의 폴리펩티드 분자 간의 서열 정보, 예로는 점수(score), 동일성(identity) 및 유사도(similarity) 등의 매개 변수(parameter)를 직접 정렬하여 결정될 수 있다(예: BLAST 2.0). 또한, 폴리뉴클레오티드 간 상동성은 상동 영역 간의 안정된 이중가닥을 이루는 조건하에서 폴리뉴클레오티드의 혼성화한 후, 단일-가닥-특이적 뉴클레아제로 분해시켜 분해된 단편의 크기를 결정함으로써 결정할 수 있다.
본 발명에서 용어, "내재적 활성"은 미생물이 천연의 상태 또는 해당 효소의 변형 전에 가지고 있는 효소의 활성 상태를 의미한다.
상기 "효소의 활성이 내재적 활성에 비해 약화된 것"은 본래 미생물이 천연의 상태 또는 변형 전의 상태에서 가지고 있는 효소의 활성과 비교하였을 때, 그 활성이 감소된 것을 의미한다. 상기 약화는 상기 효소를 코딩하는 유전자의 변이 등으로 효소 자체의 활성이 본래 미생물이 가지고 있는 효소의 활성에 비해 감소한 경우와, 이를 코딩하는 유전자의 발현 저해 또는 번역(translation) 저해 등으로 세포 내에서 전체적인 효소 활성 정도가 천연형 균주 또는 변형전의 균주에 비하여 낮은 경우, 이들의 조합 역시 포함하는 개념이다.
상기 "불활성화"는 효소를 코딩하는 유전자의 발현이 천연형 균주 또는 변형전의 균주에 비하여 전혀 발현이 되지 않는 경우 및 발현이 되더라도 그 활성이 없는 경우를 의미한다.
이러한 효소 활성의 약화 또는 불활성화는, 당해 분야에 잘 알려진 다양한 방법의 적용으로 달성될 수 있다. 상기 방법의 예로, 상기 효소의 활성이 제거된 경우를 포함하여 상기 효소의 활성이 감소되도록 돌연변이된 유전자로, 염색체상의 상기 효소를 코딩하는 유전자를 대체하는 방법; 상기 효소를 코딩하는 염색체상의 유전자의 발현 조절 서열에 변이를 도입하는 방법; 상기 효소를 코딩하는 유전자의 발현 조절 서열을 활성이 약하거나 없는 서열로 교체하는 방법; 상기 효소를 코딩하는 염색체상의 유전자의 전체 또는 일부를 결실시키는 방법; 상기 염색체상의 유전자의 전사체에 상보적으로 결합하여 상기 mRNA로부터 효소로의 번역을 저해하는 안티센스 올리고뉴클레오티드(예컨대, 안티센스 RNA)를 도입하는 방법; 상기 효소를 코딩하는 유전자의 SD 서열 앞단에 SD 서열과 상보적인 서열을 인위적으로 부가하여 2차 구조물을 형성시켜 리보솜(ribosome)의 부착이 불가능하게 만드는 법 및 해당 서열의 ORF(open reading frame)의 3' 말단에 역전사되도록 프로모터를 부가하는 RTE(Reverse transcription engineering) 방법 등이 있으며, 이들의 조합으로도 달성할 수 있으나, 상기 예에 의해 특별히 제한되는 것은 아니다.
구체적으로, 효소를 코딩하는 유전자의 일부 또는 전체를 결실하는 방법은, 세균 내 염색체 삽입용 벡터를 통해 염색체 내 내재적 목적 단백질을 암호화하는 폴리뉴클레오티드를 일부 핵산 서열이 결실된 폴리뉴클레오티드 또는 마커 유전자로 교체함으로써 수행될 수 있다. 이러한 유전자의 일부 또는 전체를 결실하는 방법의 일례로 상동 재조합에 의하여 유전자를 결실시키는 방법을 사용할 수 있다.
상기에서 "일부"란, 폴리뉴클레오티드의 종류에 따라서 상이하지만, 구체적으로는 1 내지 300개, 구체적으로는 1 내지 100개, 더욱 구체적으로는 1 내지 50개일 수 있으나, 특별히 이에 제한되는 것은 아니다.
상기에서 "상동 재조합(homologous recombination)"이란, 서로 상동성을 지닌 유전자 사슬의 좌위에서 연결 교환을 통해 일어나는 유전자 재조합을 의미한다.
구체적으로, 발현 조절 서열을 변형하는 방법은 상기 발현 조절 서열의 핵산 서열에 결실, 삽입, 비보전적 또는 보전적 치환 또는 이들의 조합으로 발현 조절 서열상의 변이를 유도하여 수행하거나, 더욱 약한 프로모터로 교체하는 등의 방법으로써 수행할 수 있다. 상기 발현 조절서열에는 프로모터, 오퍼레이터 서열, 리보좀 결합부위를 코딩하는 서열, 및 전사와 해독의 종결을 조절하는 서열을 포함한다.
아울러, 염색체상의 유전자 서열을 변형하는 방법은 상기 효소의 활성이 더욱 약화하도록 유전자 서열을 결실, 삽입, 비보전적 또는 보전적 치환 또는 이들의 조합으로 서열상의 변이를 유도하여 수행하거나, 더욱 약한 활성을 갖도록 개량된 유전자 서열 또는 활성이 없도록 개량된 유전자 서열로 교체함으로써 수행할 수 있다.
본 발명의 일 구현예에서는, 상기 활성의 약화 또는 불활성화는 상기 6-포스포글루코네이트 탈수소효소를 코딩하는 유전자 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제를 코딩하는 유전자 내로, 하나 이상의 염기쌍의 삽입에 의한 삽입 돌연변이, 상기 유전자 내의 하나 이상의 염기쌍의 결실을 갖는 결실 돌연변이, 및 상기 유전자 내의 넌센스 코돈 또는 상이 코돈을 도입시키는 염기쌍의 전이(transition) 또는 전환(transversion) 돌연변이로 이루어지는 군으로부터 선택되는 하나 이상의 돌연변이 방법을 통해 수행할 수 있음을 확인하였다.
본 발명에서 상기 에스케리키아속(Escherichia) 미생물은 구체적으로 대장균(Escherichia coli)일 수 있으나, 이에 한정되지 않는다.
구체적으로 본 발명에서 상기 edd 및 eda의 활성의 약화 또는 불활성화에 의하여 L-트립토판 생산능을 갖는 에스케리키아속 미생물의 모균주는 트립토판 생산능을 가지는 에스케리키아속 미생물이라면 특별히 제한되지 않는다. 트립토판 생산능을 가지는 미생물은, 그 예로, 트립토판 생합성 경로를 강화하기 위하여, 경쟁경로의 유전자, 트립토판 오페론의 방향성 경로의 조절자, 트립토판 유입유전자, 트립토판 유입 및 분해 유전자의 활성의 약화 또는 불활성화 시키거나, 및/또는 트립토판 오페론의 활성을 과발현시킨 미생물일 수 있다. 상기 활성의 약화 또는 불활성화 방법은 상기에서 설명한 바와 같으며, 공지의 방법은 제한 없이 포함된다. 또한, 트립토판 오페론 활성의 과발현은 공지의 방법은 제한없이 포함되나, 그 예로 오페론 유전자의 염기서열 일부 또는 전부 그 자체 혹은 외부로부터 도입된 발현조절부위를 포함하는 폴리뉴클레오타이드를 추가로 염색체에 삽입하는 방법, 벡터 시스템에 도입하는 방법에 의하여 카피수를 증가시키는 방법, 유전자의 발현을 조절하는 발현조절부위를 다른 조절서열로 치환, 발현조절부위의 염기서열 전체 혹은 일부 돌연변이가 유발된 변형, 유전자 자체의 변이도입에 의한 오페론 활성 강화 등에 의한 방법 등을 포함할 수 있으나, 이에 한정되는 것은 아니다. 구체적으로는 pheA, trpR, mtr 및 tnaAB 유전자의 전체 또는 일부가 결실, 및/또는 트립토판 오페론이 과발현된 대장균일 수 있다.
본 발명에서, edd, eda, pheA, trpR, mtr, tnaAB 유전자 및 트립토판 오페론 및 이들이 코딩하는 단백질 서열을 비롯하여, 본 발명에서 사용되는 유전자 및 단백질 서열들은 공지의 데이터 베이스에서 얻을 수 있으며, 그 예로 NCBI의 GenBank 등에서 얻을 수 있으나, 이에 제한되지 않는다. 또한, pheA, trpR, mtr 및 tnaAB 유전자에 대한 구체적인 내용은 한국 특허등록 제10-0792095호에 기재되어 있는 내용을 참고할 수 있으며, 상기 특허의 명세서 전체는 본 발명의 참고자료로서 포함될 수 있다.
본 발명의 구체적인 실시예들에 비추어, 다양한 모균주에서 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 불활성화시킨 결과, Entner-Doudoroff 경로를 갖고 있는 에스케리키아속 미생물이라면 모균주의 종류에 상관없이 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 함께 약화시키거나 불활성화시키는 경우에는 L-트립토판의 생산능이 유의적으로 향상되는 것으로 확인되었다.
본 발명의 또 다른 일 양태에 따르면, 본 발명의 내재적 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성이 약화 또는 불활성화되어, L-트립토판 생산능을 갖는 에스케리키아속 미생물을 배양하는 단계; 및 상기 미생물의 배양 단계에 따른 배양물 또는 상기 미생물로부터 L-트립토판을 회수하는 단계를 포함하는, L-트립토판의 제조 방법을 제공한다.
본 발명의 미생물의 배양에 사용되는 배지 및 기타 배양 조건은 통상의 에스케리키아속 미생물의 배양에 사용되는 배지라면 특별한 제한없이 어느 것이나 사용할 수 있으나, 구체적으로는 본 발명의 미생물을 적당한 탄소원, 질소원, 인원, 무기화합물, 아미노산 및/또는 비타민 등을 함유한 통상의 배지 내에서 호기성 조건 하에서 온도, pH 등을 조절하면서 배양할 수 있다.
본 발명에서 상기 탄소원으로는 글루코오스, 프룩토오스, 수크로오스, 말토오스, 만니톨, 소르비톨 등과 같은 탄수화물; 당 알코올, 글리세롤, 피루브산, 락트산, 시트르산 등과 같은 알코올; 유기산, 글루탐산, 메티오닌, 리신 등과 같은 아미노산 등이 포함될 수 있다. 또한, 전분 가수분해물, 당밀, 블랙스트랩 당밀, 쌀겨울, 카사버, 사탕수수 찌꺼기 및 옥수수 침지액 같은 천연의 유기 영양원을 사용할 수 있으며, 구체적으로는 글루코오스 및 살균된 전처리 당밀(즉, 환원당으로 전환된 당밀) 등과 같은 탄수화물이 사용될 수 있으며, 그 외의 적정량의 탄소원을 제한 없이 다양하게 이용할 수 있다. 이들 탄소원은 단독으로 사용되거나 2 종 이상이 조합되어 사용될 수 있다.
상기 질소원으로는 암모니아, 황산암모늄, 염화암모늄, 초산암모늄, 인산암모늄, 탄산안모늄, 질산암모늄 등과 같은 무기질소원; 글루탐산, 메티오닌, 글루타민 등과 같은 아미노산, 펩톤, NZ-아민, 육류 추출물, 효모 추출물, 맥아 추출물, 옥수수 침지액, 카세인 가수분해물, 어류 또는 그의 분해생성물, 탈지 대두 케이크 또는 그의 분해 생성물 등과 같은 유기 질소원이 사용될 수 있다. 이들 질소원은 단독으로 사용되거나 2 종 이상이 조합되어 사용될 수 있다.
상기 인원으로는 인산 제1칼륨, 인산 제2칼륨, 또는 이에 대응되는 소디움-함유 염 등이 포함될 수 있다. 무기화합물로는 염화나트륨, 염화칼슘, 염화철, 황산마그네슘, 황산철, 황산망간, 탄산칼슘 등이 사용될 수 있으며, 그 외에 아미노산, 비타민 및/또는 적절한 전구체 등이 포함될 수 있다. 이들 배지 또는 전구체는 배양물에 회분식 또는 연속식으로 첨가될 수 있다.
본 발명에서, 미생물의 배양 중에 수산화암모늄, 수산화칼륨, 암모니아, 인산, 황산 등과 같은 화합물을 배양물에 적절한 방식으로 첨가하여, 배양물의 pH를 조정할 수 있다. 또한, 배양 중에는 지방산 폴리글리콜 에스테르와 같은 소포제를 사용하여 기포 생성을 억제할 수 있다. 또한, 배양물의 호기 상태를 유지하기 위하여, 배양물 내로 산소 또는 산소 함유 기체를 주입하거나 혐기 및 미호기 상태를 유지하기 위해 기체의 주입 없이 혹은 질소, 수소 또는 이산화탄소 가스를 주입할 수 있다.
배양물의 온도는 구체적으로는 27℃ 내지 40℃보다 구체적으로는 30℃ 내지 37℃일 수 있으나 이에 제한되지 않는다. 배양 기간은 유용 물질의 원하는 생성량이 수득될 때까지 계속될 수 있으며, 구체적으로는 10 시간 내지 100 시간일 수 있으나 이에 제한되지 않는다.
상기 L-트립토판을 회수하는 단계는 본 발명의 미생물의 배양 방법, 예를 들어 회분식, 연속식 또는 유가식 배양 방법 등에 따라 당해 기술 분야에 공지된 적합한 방법을 이용하여 배양액으로부터 목적하는 L-트립토판을 회수할 수 있다.
상기 회수 단계는 정제 공정을 포함할 수 있다.
본 발명은 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제의 활성을 약화 또는 불활성화시킴으로써 L-트립토판 생산능을 갖는 에스케리키아속 미생물을 제공하여, 이를 이용해 L-트립토판을 보다 고수율로 효율적이고 경제적으로 제조할 수 있는 효과를 나타낸다.
이하, 실시예들을 통하여 본 발명을 보다 상세히 설명한다. 다만, 이들 실시예는 본 발명을 예시적으로 설명하기 위한 것에 불과하므로 본 발명의 범위가 이들 실시예에 의해 한정되는 것으로 해석되어서는 아니된다.
비교예
1: Δ
pheA
Δ
trpR
Δ
mtr
Δ
tnaAB
/
pCL1920
-
Ptrc
-
trpO
인
모균주의
제작
(1)
pheA
,
trpR
,
mtr
및
tnaAB
유전자에 의하여 코딩되는 단백질이 불활성화된 야생형 균주의 제작
모균주의 트립토판 생합성경로를 강화하기 위해 경쟁경로의 유전자인 pheA, 트립토판 오페론과 방향성 경로의 조절자인 trpR, 트립토판 유입유전자인 mtr, 트립토판 유입 및 분해유전자인 tnaA 및 tnaB 를 모두 불활성화시켜 본 발명의 트립토판 생산능을 더욱 더 잘 확인할 수 있도록 하였다. 대장균 W3110 (ATCC®39936TM)에서 pheA 유전자에 의해 코딩되는 chorismate mutase/prephenate hydratase; trpR 유전자에 의해 코딩되는 trpR transcriptional repressor; mtr 유전자에 의해 코딩되는 tryptophan/Indole:H+ symporter; tnaA 및 tnaB 유전자에 의해 코딩되는 오페론 형태의 tryptophanase 및 trpytpophan:H+symporter를 유전자 상동 재조합에 의하여 모두 불활성화시켰다. 이를 위하여, Datsenko KA 등이 개발한 람다 레드 재조합 효소(lambda Red recombinase)를 이용한 1단계 불활성화(one step inactivation) 방법을 사용하였으며(One-step inactivation of chromosomal genes in Escherichia coli K-12 using PCR products, Datsenko KA, Wanner BL., Proc Natl Acad Sci USA, 2000, Jun 6;97(12):6640-5), 대한민국등록특허 제10-0792095호에 기반하여 불활성화를 진행하였다. 본 비교예 1을 포함하여, 후술하는 비교예 2 및 실시예 1, 3에 사용된 프라이머의 서열은 하기 표 3에 나타내었다.
구체적으로, 서열번호 6의 서열을 갖는 pheA 유전자의 일부분과 pKD3 유전자의 클로람페니콜(Chloramphenicol) 내성 유전자의 일부 염기서열을 갖는 프라이머 1 및 2를 이용하여 pKD3를 주형으로 하여 PCR법에 의해 약 1,100 쌍의 유전자 단편을 증폭하였다. 그 다음, 상기 PCR 증폭을 통하여 얻어진 DNA 단편을 0.8% 아가로스 겔에서 전기 영동한 후 용리하여 2차 PCR의 주형으로 사용하였다. 그 다음, 대장균 pheA 유전자의 5' 및 3' DNA 단편을 얻기 위하여, 프라이머 3과 4, 프라이머 5와 6을 이용하여 대장균 W3110의 염색체를 주형으로 하여 PCR법에 의해 약 250 쌍의 유전자 단편을 각각 증폭하였다. 그 다음, 상기 PCR 증폭을 통하여 얻어진 DNA 단편을 0.8% 아가로스 겔에서 전기 영동한 후 용리하여 2차 PCR의 주형으로 사용하였다.
상기에서, 프라이머 1과 4의 18쌍의 염기서열이 상보적이며, 프라이머 2와 5의 20 쌍의 염기서열이 상보적이기 때문에, 프라이머 1과 2를 이용하여 얻어진 단편, 프라이머 3과 4로 얻어진 단편, 프라이머 5와 6으로 얻어진 단편들은 하나의 단편으로 연결될 수 있다. 상기 얻어진 PCR 단편들을 프라이머 없이 5회 PCR법으로 증폭한 다음, 프라이머 3과 6를 첨가한 후 다시 25회 PCR법으로 증폭하였다. 그 결과 약 1,600 염기쌍 크기의 유전자 단편이 증폭되었다.
그 다음, Datsenko KA 등이 개발한 방법에 따라 pKD46으로 형질 전환된 대장균 W3110을 컴피턴트 세포로 제작한 후, PCR로 얻어진 1,600 염기쌍 크기의 유전자 단편을 유입하여 클로람페니콜이 함유된(30 ㎎/L) LB 고체 배지에 도말하였다. 얻어진 균주는 다시 프라이머 3과 6을 이용한 PCR법으로 얻어진 크기가 1,600 염기쌍으로 pheA가 불활성화되었다는 것을 확인하고, W3110 ΔpheA 균주를 제작하였다.
위와 같은 방법으로 서열번호 8의 서열을 갖는 trpR, 서열번호 10의 서열을 갖는 mtr, 서열번호 12와 14의 서열을 갖는 tnaAB 유전자들에 의해 코딩되는 단백질들을 하기 표 3의 프라이머를 이용하여 불활성화시켜, W3110 ΔpheAΔtrpRΔmtrΔtnaAB 균주를 제작하였다.
(2) 트립토판
생산능을
나타내는 유전자들이 도입된 벡터의 제작
상기 제조된 W3110ΔpheAΔtrpRΔmtrΔtnaAB 야생형 균주에 트립토판 생산능을 부여하기 위하여 pCL1920 벡터에 Ptrc 프로모터 및 트립토판 오페론 유전자를 삽입하여 재조합 벡터 pCL1920-Ptrc-trpO를 제작하였다.
구체적으로는, Ptrc 프로모터를 pCL1920 벡터로 삽입하기 위하여 pCL1920의 플라스미드를 회수하여 제한효소 HindⅢ 및 PstI로 처리하여 준비하고, Ptrc 프로모터는 pTrcHis B (invitrogen, USA) 벡터를 주형으로 하고, 프라이머 23과 24를 이용해 94℃에서 30초간 변성, 58℃에서 30초간 어닐링, 72℃에서 30초간 중합하는 조건을 30회 반복하는 PCR법에 의해 준비하였다. 상기 준비된 Ptrc 프로모터 단편은 제한효소 HindⅢ 및 PstI로 절단한 후, pCL1920과 라이게이션하여 pCL1920-Ptrc 벡터를 제작하였다.
그 다음, pCL1920_Ptrc_trpO 벡터를 제작하기 위하여 pCL1920_Ptrc 벡터는 PstI 및 알칼리 포스파타아제(Alkaline phosphatase) 처리하여 준비하였고, 트립토판 오페론 유전자는 대장균 KCCM10812P(대한민국등록특허 제10-0792095호)의 염색체 DNA로부터 증폭하였다. 해당 오페론 유전자의 첫 번째 유전자인 trpE 유전자는 피드백 저항성 형태를 갖는다. 증폭을 위하여 대장균 KCCM10812P의 염색체 DNA를 주형으로 프라이머 25와 26을 이용해 94℃에서 30초간 변성, 58℃에서 30초간 어닐링, 72℃에서 5분간 중합하는 조건을 30회 반복하는 PCR법을 수행하였다. 상기 얻어진 DNA 단편은 PstI로 처리하여 준비된 pCL1920_Ptrc 벡터와 라이게이션하였고, 그 결과 얻어진 벡터를 pCL1920_Ptrc_trpO(서열번호 15)로 명명하였다.
(3) 트립토판
오페론을
포함하는 벡터가 삽입된 균주의 제작
비교예 1의 (1)에서 제작된 균주를 컴피턴트 세포로 제작한 후, 비교예1의 (2)에서 만든 벡터를 도입하여 트립토판 생산능을 가지는 W3110ΔpheAΔtrpRΔmtrΔtnaAB/pCL1920-Ptrc-trpO 야생형 균주를 제작하였다.
실시예
1:
비교예
1의
모균주로부터
Δ
edd
Δ
eda
인 균주의 제작
상기 비교예 1에서 제작된 W3110 ΔpheAΔtrpRΔmtrΔtnaAB/pCL1920-Ptrc-trpO 야생형 균주에 있어서, edd 및 eda 유전자군을 상동 재조합에 의하여 동시에 결실시킴으로써 edd(서열번호 2) 및 eda 유전자(서열번호 4)에 의해 코딩되는 6-포스포글루코네이트 탈수소효소 및 2-케토-3-데옥시-6-포스포글루코네이트가 모두 불활성화된 균주를 제작하였다.
구체적으로, 상기 균주를 제작하기 위하여, Datsenko KA 등이 개발한 람다 레드 재조합 효소를 이용한 돌연변이 제작 기법인 1단계 불활성화 방법을 사용하였다. 유전자 내부로의 삽입을 확인하기 위한 마커로는 pUCprmfmloxC의 클로람페니콜 유전자를 사용하였다(대한민국공개특허 2009-007554). 상기 edd-eda 유전자군의 일부분과 pUCprmfmloxC 유전자의 클로람페니콜 내성 유전자의 일부 염기서열을 갖는 프라이머 27과 프라이머 28을 이용하여 pUCprmfmloxC를 주형으로 하고 94℃에서 30초간 변성, 55℃에서 30초간 어닐링, 72℃에서 1분간 중합하는 조건을 30회 반복하는 PCR법에 의해 약 1,200쌍의 유전자 단편을 증폭하였다.
또한, PCR 증폭을 통하여 얻어진 DNA 단편은 0.8% agarose겔 전기영동 후 용리하여 2차 PCR의 주형으로 사용하였다. 2차 PCR은 1차 DNA 단편의 5' 및 3' 영역의 20쌍의 상보적 염기서열을 갖도록 하였고, edd-eda 유전자군의 5' 및 3' 영역을 더한 프라이머 29과 프라이머 30을 이용하여 용리된 1차 PCR 산물을 주형으로 94℃에서 30초간 변성, 55℃에서 30초간 어닐링, 72℃에서 1분간 중합하는 조건을 30회 반복하는 PCR법에 의해 약 1,300쌍의 유전자 단편을 증폭하였다. 상기 과정을 거쳐 얻어진 DNA단편은 0.8% 아가로스 겔 전기영동 후 용리하여 재조합에 사용하였다.
Datsenko KA 등이 개발한 방법에 따라 pKD46으로 형질 전환된 대상 대장균을 컴피턴트한 상태로 제조 후, PCR법으로 얻어진 1,300 염기쌍 크기의 유전자 단편을 유입하여 형질전환시켰다. 얻어진 균주는 클로람페니콜 내성을 갖는 LB에서 선별하였으며, 다시 프라이머 31과 프라이머 32를 이용한 PCR로 얻어진 크기가 1,626 염기쌍으로 edd-eda 유전자군이 결실되었다는 것을 확인하였다.
클로람페니콜 내성을 가진 1차 재조합 균주는 pKD46을 제거한 후, pJW168을 도입하여 클로람페니콜 마커 유전자를 균체로부터 제거하였다(Gene, (2000) 247, 255-64). 최종적으로 얻어진 균체는 프라이머 31과 32로 PCR을 통해 얻어진 580쌍의 증폭 산물로 의도한 대로 결실이 이루어졌음을 확인하였다. 그리고 이 균주를 컴피턴트한 상태로 제조 후, 비교예 1에서 제작된 벡터를 유입하여 형질전환시켜 트립토판 생산능을 가지는 균주 W3110ΔpheAΔtrpRΔmtrΔtnaABΔeddΔeda/pCL1920-Ptrc-trpO을 제작하였다.
실시예
2: Δ
edd
Δ
eda
균주의 트립토판
생산능
확인
상기 비교예 1 및 실시예 1에서 제작된 균주를 이용하여 역가 평가를 진행하였다. 역가 평가를 위하여 균체를 백금이로 접종한 후, LB 고체 배지에서 밤새 배양하고, 하기 표 1의 조성을 갖는 25 ㎖의 플라스크 역가 배지에 한 백금이 씩 접종하였다. 균주 접종 후 37℃, 200 rpm에서 42시간 동안 배양하고, 그로부터 얻어진 결과를 하기 표 2에 나타내었다. 모든 결과는 3개의 플라스크 결과의 평균값을 사용하였다.
조성물 | 농도(리터당) |
글루코스 | 30 g |
K2HPO4 | 1 g |
(NH4)2SO4 | 10 g |
NaCl | 1 g |
MgSO4ㆍ7H2O | 1 g |
구연산나트륨 | 5 g |
효모액기스 | 2 g |
탄산칼슘 | 40 g |
구연산나트륨 | 5 g |
페닐알라닌 | 0.15 g |
타이로신 | 0.1 g |
pH | 6.8 g |
균주 | OD | 소모당 (g/L)* |
L-트립토판 (g/L)** |
W3110ΔpheAΔtrpRΔmtr ΔtnaAB/pCL1920-Ptrc-trpO | 30.1 | 24.7 | 0.78 |
실시예 1 (W3110ΔpheAΔtrpRΔmtrΔtnaABΔeddΔeda/pCL1920-Ptrc-trpO) |
29.5 | 25.1 | 1.03 |
* 22시간 측정치
** 42시간 측정치
상기 실험 결과, 상기 표 2에 나타난 바와 같이 본 발명에서 제시한 edd-eda 유전자군의 결실시, 당 소모 속도는 모균주인 비교예 1과 유의차가 없었으나, 트립토판 생산량은 모균주 대비 약 32% 이상 향상된 것으로 확인되었다. 이 결과는 Enter Doudoroff 경로의 결실에 의해 기질인 6-포스포글루코네이트의 손실없이 루불로오스 5-포스페이트(Rubulose 5-phosphate)까지 반응이 이루어졌기 때문에 NADPH는 물론, PRPP (5-phosphoribosyl-1-pyrophosphate) 및 E4P (Erythrose 4-phosphate)의 양이 증가하여 결과적으로 트립토판 생산능력이 증가된 것으로 여겨진다.
실시예
3:
기탁된
모균주로부터
Δ
edd
Δ
eda
인 균주의 제작
한국미생물보존센터에 기탁된 L-트립토판 생산 대장균 KCCM11166P(대한민국등록특허 제10-1261147호)를 상기 실시예 1과 동일한 방법으로 처리하여, edd-eda 유전자군이 결실된 KCCM11166PΔeddΔeda 균주를 제작하였다.
유전자 | 프라이머 번호 |
서열 (5'→3') | 서열번호 |
pheA | 1 | AGGCAACACTATGACATCGTGTAGGCTGGAGCTGCTTC | 16 |
2 | GGTCGCCATTAACAACGTGGCATATGAATATCCTCCTTAG | 17 | |
3 | TATTGAGTGTATCGCCAAC | 18 | |
4 | CGATGTCATAGTGTTGCC | 19 | |
5 | CCACGTTGTTAATGGCGACC | 20 | |
6 | TTCATTGAACGGGTGATTTC | 21 | |
trpR | 7 | TCCGCACGTTTATGATATGCTATCGTACTCTTTAGCGAGTACAACCGGGGGTGTAGGCTGGAGCTGCTTC | 22 |
8 | GCCACGTCTTATCAGGCCTACAAAATCAATCGCTTTTCAGCAACACCTCTCATATGAATATCCTCCTTAG | 23 | |
9 | GCGCCGGGCGTATCGACGCA | 24 | |
10 | GCATATCATAAACGTGCGGA | 25 | |
11 | TGTAGGCCTGATAAGACGTG | 26 | |
12 | AAGGGGCGATCGGCGTGTTT | 27 | |
mtr | 13 | ATGGCAACACTAACCACCACCCAAACGTCACCGTCGCTGCTTGGCGGCGTGTGTAGGCTGGAGCTGCTTC | 28 |
14 | TTACTGATACACCGGCAGTAAATTAAAGCTCGATAAAATATGCACCAGTGCATATGAATATCCTCCTTAG | 29 | |
15 | GCAGCCGTTACATTGGTAAC | 30 | |
16 | GTGGTGGTTAGTGTTGCCAT | 31 | |
17 | TACTGCCGGTGTATCAGTAA | 32 | |
18 | TCAAACCGTCAGCACGGCTG | 33 | |
tnaAB |
19 | ATGAAGGATTATGTAATGGAAAACTTTAAACATCTCCCTGAACCGTTCCGGTGTAGGCTGGAGCTGCTTC | 34 |
20 | TTAGCCAAATTTAGGTAACACGTTAAAGACGTTGCCGAACCAGCACAAAACATATGAATATCCTCCTTAG | 35 | |
21 | TTAAGCGAAATCACCGGGGAA | 36 | |
22 | ATGTCCGAGCACTGGCGC | 37 | |
pCL1920-Ptrc-trpO | 23 | CCCAAGCTTGCTGTTGACAATTAATCAT | 38 |
24 | AAAACTGCAGCTGTTTCCTGTGTGAAAT | 39 | |
25 | AAAACTGCAGATGCAAACACAAAAACCGACT | 40 | |
26 | AAAACTGCAGTTAACTGCGCGTCGCCGCTTTC | 41 | |
Edd-eda | 27 | AAACGCGTTGTGAATCATCCTGCTCTGACAACTCAATTTCAGGAGCCTTTGCCGCCAGCTGAAGCTTTAC | 42 |
28 | ACAGCACGCTTTTCAGCGCCAGGTAGTCACGGTAGTTAGCCGGAGAAATATAGTGGATCTGATGGGTACC | 43 | |
29 | TGCCCTATGAGCTCCGGTTACAGGCGTTTCAGTCATAAATCCTCTGAATGAAACGCGTTGTGAATCATCC | 44 | |
30 | ATCGCCCGCTTCCAGCGCATCTGCCGGAACCAGCCAGGAACCACCGATGCACAGCACGCTTTTCAGCGCC | 45 | |
31 | CATGATCTTGCGCAGATTGTA | 46 | |
32 | CATGATCTTGCGCAGATTGTA | 47 |
실시예
4: Δ
edd
Δ
eda
균주의 트립토판
생산능
확인
상기 기탁균주 KCCM11166P 및 실시예 3에서 제작된 균주를 이용하여 역가 평가를 진행하였다.
역가 평가를 위하여 균체를 백금이로 접종한 후, LB 고체 배지에서 밤새 배양하고, 하기 표 4의 조성을 갖는 25 ㎖의 플라스크 역가 배지에 한 백금이 씩 접종하였다. 균주 접종 후 37℃, 200 rpm에서 42시간 동안 배양하고, 그로부터 얻어진 결과를 하기 표 5에 나타내었다. 모든 결과는 3개의 플라스크 결과의 평균값을 사용하였다.
조성물 | 농도(리터당) |
글루코스 | 60 g |
K2HPO4 | 1 g |
(NH4)2SO4 | 10 g |
NaCl | 1 g |
MgSO4ㆍ7H2O | 1 g |
구연산나트륨 | 5 g |
효모액기스 | 2 g |
탄산칼슘 | 40 g |
구연산나트륨 | 5 g |
페닐알라닌 | 0.15 g |
타이로신 | 0.1 g |
pH | 6.8 g |
균주 | OD | 소모당 (g/L)* |
L-트립토판 (g/L)** |
KCCM11166P | 22.9 | 31.0 | 5.7 |
KCCM11166PΔeddΔeda | 23.6 | 30.7 | 6.9 |
* 33시간 측정치
** 48간 측정치
상기 실험 결과, 상기 표 5에 나타난 바와 같이 본 발명에서 제시한 edd-eda 유전자군의 결실시, 당 소모 속도는 모균주와 유의한 차이가 없었으나, 트립토판 생산량은 모균주 대비 약 21% 이상 향상된 것으로 확인되었다. 이 결과는 상기 실시예 2에서 언급한 것과 같은 이유로 트립토판 생산능력이 증가된 것이라고 할 수 있다.
본 발명자들은 KCCM11166P 균주 기반, edd-eda 유전자군의 불활성화 균주에서 트립토판 생산능력이 증가함을 확인하고, 상기 균주를 "CA04-2800"으로 명명한 후 부다페스트 조약 하에 국제기탁기관인 한국미생물보존센터 (KCCM)에 2013년 11월 15일에 기탁하여 수탁번호 KCCM11473P를 부여받았다.
상기와 같은 결과들은 Entner-Doudoroff 경로를 갖고 있는 에스케리키아속 미생물에 있어서, edd-eda 활성을 동시에 불활성화시킬 경우, 불활성화시키지 않은 미생물에 비해 트립토판 생산능이 증가되는 것을 시사하는 것이다.
본 명세서는 본 발명의 기술 분야에서 통상의 지식을 가진 자이면 충분히 인식하고 유추할 수 있는 내용은 그 상세한 기재를 생략하였으며, 본 명세서에 기재된 구체적인 예시들 이외에 본 발명의 기술적 사상이나 필수적 구성을 변경하지 않는 범위내에서 보다 다양한 변형이 가능하다. 따라서 본 발명은 본 명세서에서 구체적으로 설명하고 예시한 것과 다른 방식으로 실시될 수 있으며, 이는 본 발명의 기술 분야에 통상의 지식을 가진 자이면 이해할 수 있는 사항이다.
<110> CJ CheilJedang Corporation
<120> A MICROORGANISM OF ESCHERICHIA GENUS HAVING L-TRYPTOPHAN
PRODUCING ACTIVITY AND METHOD FOR PRODUCING L-TRYPTOPHAN USING
THE SAME
<130> PA131256/KR
<160> 47
<170> KopatentIn 2.0
<210> 1
<211> 603
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(603)
<223> edd
<400> 1
Met Asn Pro Gln Leu Leu Arg Val Thr Asn Arg Ile Ile Glu Arg Ser
1 5 10 15
Arg Glu Thr Arg Ser Ala Tyr Leu Ala Arg Ile Glu Gln Ala Lys Thr
20 25 30
Ser Thr Val His Arg Ser Gln Leu Ala Cys Gly Asn Leu Ala His Gly
35 40 45
Phe Ala Ala Cys Gln Pro Glu Asp Lys Ala Ser Leu Lys Ser Met Leu
50 55 60
Arg Asn Asn Ile Ala Ile Ile Thr Ser Tyr Asn Asp Met Leu Ser Ala
65 70 75 80
His Gln Pro Tyr Glu His Tyr Pro Glu Ile Ile Arg Lys Ala Leu His
85 90 95
Glu Ala Asn Ala Val Gly Gln Val Ala Gly Gly Val Pro Ala Met Cys
100 105 110
Asp Gly Val Thr Gln Gly Gln Asp Gly Met Glu Leu Ser Leu Leu Ser
115 120 125
Arg Glu Val Ile Ala Met Ser Ala Ala Val Gly Leu Ser His Asn Met
130 135 140
Phe Asp Gly Ala Leu Phe Leu Gly Val Cys Asp Lys Ile Val Pro Gly
145 150 155 160
Leu Thr Met Ala Ala Leu Ser Phe Gly His Leu Pro Ala Val Phe Val
165 170 175
Pro Ser Gly Pro Met Ala Ser Gly Leu Pro Asn Lys Glu Lys Val Arg
180 185 190
Ile Arg Gln Leu Tyr Ala Glu Gly Lys Val Asp Arg Met Ala Leu Leu
195 200 205
Glu Ser Glu Ala Ala Ser Tyr His Ala Pro Gly Thr Cys Thr Phe Tyr
210 215 220
Gly Thr Ala Asn Thr Asn Gln Met Val Val Glu Phe Met Gly Met Gln
225 230 235 240
Leu Pro Gly Ser Ser Phe Val His Pro Asp Ser Pro Leu Arg Asp Ala
245 250 255
Leu Thr Ala Ala Ala Ala Arg Gln Val Thr Arg Met Thr Gly Asn Gly
260 265 270
Asn Glu Trp Met Pro Ile Gly Lys Met Ile Asp Glu Lys Val Val Val
275 280 285
Asn Gly Ile Val Ala Leu Leu Ala Thr Gly Gly Ser Thr Asn His Thr
290 295 300
Met His Leu Val Ala Met Ala Arg Ala Ala Gly Ile Gln Ile Asn Trp
305 310 315 320
Asp Asp Phe Ser Asp Leu Ser Asp Val Val Pro Leu Met Ala Arg Leu
325 330 335
Tyr Pro Asn Gly Pro Ala Asp Ile Asn His Phe Gln Ala Ala Gly Gly
340 345 350
Val Pro Val Leu Val Arg Glu Leu Leu Lys Ala Gly Leu Leu His Glu
355 360 365
Asp Val Asn Thr Val Ala Gly Phe Gly Leu Ser Arg Tyr Thr Leu Glu
370 375 380
Pro Trp Leu Asn Asn Gly Glu Leu Asp Trp Arg Glu Gly Ala Glu Lys
385 390 395 400
Ser Leu Asp Ser Asn Val Ile Ala Ser Phe Glu Gln Pro Phe Ser His
405 410 415
His Gly Gly Thr Lys Val Leu Ser Gly Asn Leu Gly Arg Ala Val Met
420 425 430
Lys Thr Ser Ala Val Pro Val Glu Asn Gln Val Ile Glu Ala Pro Ala
435 440 445
Val Val Phe Glu Ser Gln His Asp Val Met Pro Ala Phe Glu Ala Gly
450 455 460
Leu Leu Asp Arg Asp Cys Val Val Val Val Arg His Gln Gly Pro Lys
465 470 475 480
Ala Asn Gly Met Pro Glu Leu His Lys Leu Met Pro Pro Leu Gly Val
485 490 495
Leu Leu Asp Arg Cys Phe Lys Ile Ala Leu Val Thr Asp Gly Arg Leu
500 505 510
Ser Gly Ala Ser Gly Lys Val Pro Ser Ala Ile His Val Thr Pro Glu
515 520 525
Ala Tyr Asp Gly Gly Leu Leu Ala Lys Val Arg Asp Gly Asp Ile Ile
530 535 540
Arg Val Asn Gly Gln Thr Gly Glu Leu Thr Leu Leu Val Asp Glu Ala
545 550 555 560
Glu Leu Ala Ala Arg Glu Pro His Ile Pro Asp Leu Ser Ala Ser Arg
565 570 575
Val Gly Thr Gly Arg Glu Leu Phe Ser Ala Leu Arg Glu Lys Leu Ser
580 585 590
Gly Ala Glu Gln Gly Ala Thr Cys Ile Thr Phe
595 600
<210> 2
<211> 1812
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(1812)
<223> edd
<400> 2
atgaatccac aattgttacg cgtaacaaat cgaatcattg aacgttcgcg cgagactcgc 60
tctgcttatc tcgcccggat agaacaagcg aaaacttcga ccgttcatcg ttcgcagttg 120
gcatgcggta acctggcaca cggtttcgct gcctgccagc cagaagacaa agcctctttg 180
aaaagcatgt tgcgtaacaa tatcgccatc atcacctcct ataacgacat gctctccgcg 240
caccagcctt atgaacacta tccagaaatc attcgtaaag ccctgcatga agcgaatgcg 300
gttggtcagg ttgcgggcgg tgttccggcg atgtgtgatg gtgtcaccca ggggcaggat 360
ggaatggaat tgtcgctgct aagccgcgaa gtgatagcga tgtctgcggc ggtggggctg 420
tcccataaca tgtttgatgg tgctctgttc ctcggtgtgt gcgacaagat tgtcccgggt 480
ctgacgatgg cagccctgtc gtttggtcat ttgcctgcgg tgtttgtgcc gtctggaccg 540
atggcaagcg gtttgccaaa taaagaaaaa gtgcgtattc gccagcttta tgccgaaggt 600
aaagtggacc gcatggcctt actggagtca gaagccgcgt cttaccatgc gccgggaaca 660
tgtactttct acggtactgc caacaccaac cagatggtgg tggagtttat ggggatgcag 720
ttgccaggct cttcttttgt tcatccggat tctccgctgc gcgatgcttt gaccgccgca 780
gctgcgcgtc aggttacacg catgaccggt aatggtaatg aatggatgcc gatcggtaag 840
atgatcgatg agaaagtggt ggtgaacggt atcgttgcac tgctggcgac cggtggttcc 900
actaaccaca ccatgcacct ggtggcgatg gcgcgcgcgg ccggtattca gattaactgg 960
gatgacttct ctgacctttc tgatgttgta ccgctgatgg cacgtctcta cccgaacggt 1020
ccggccgata ttaaccactt ccaggcggca ggtggcgtac cggttctggt gcgtgaactg 1080
ctcaaagcag gcctgctgca tgaagatgtc aatacggtgg caggttttgg tctgtctcgt 1140
tatacccttg aaccatggct gaataatggt gaactggact ggcgggaagg ggcggaaaaa 1200
tcactcgaca gcaatgtgat cgcttccttc gaacaacctt tctctcatca tggtgggaca 1260
aaagtgttaa gcggtaacct gggccgtgcg gttatgaaaa cctctgccgt gccggttgag 1320
aaccaggtga ttgaagcgcc agcggttgtt tttgaaagcc agcatgacgt tatgccggcc 1380
tttgaagcgg gtttgctgga ccgcgattgt gtcgttgttg tccgtcatca ggggccaaaa 1440
gcgaacggaa tgccagaatt acataaactc atgccgccac ttggtgtatt attggaccgg 1500
tgtttcaaaa ttgcgttagt taccgatgga cgactctccg gcgcttcagg taaagtgccg 1560
tcagctatcc acgtaacacc agaagcctac gatggcgggc tgctggcaaa agtgcgcgac 1620
ggggacatca ttcgtgtgaa tggacagaca ggcgaactga cgctgctggt agacgaagcg 1680
gaactggctg ctcgcgaacc gcacattcct gacctgagcg cgtcacgcgt gggaacagga 1740
cgtgaattat tcagcgcctt gcgtgaaaaa ctgtccggtg ccgaacaggg cgcaacctgt 1800
atcacttttt aa 1812
<210> 3
<211> 213
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(213)
<223> eda
<400> 3
Met Lys Asn Trp Lys Thr Ser Ala Glu Ser Ile Leu Thr Thr Gly Pro
1 5 10 15
Val Val Pro Val Ile Val Val Lys Lys Leu Glu His Ala Val Pro Met
20 25 30
Ala Lys Ala Leu Val Ala Gly Gly Val Arg Val Leu Glu Val Thr Leu
35 40 45
Arg Thr Glu Cys Ala Val Asp Ala Ile Arg Ala Ile Ala Lys Glu Val
50 55 60
Pro Glu Ala Ile Val Gly Ala Gly Thr Val Leu Asn Pro Gln Gln Leu
65 70 75 80
Ala Glu Val Thr Glu Ala Gly Ala Gln Phe Ala Ile Ser Pro Gly Leu
85 90 95
Thr Glu Pro Leu Leu Lys Ala Ala Thr Glu Gly Thr Ile Pro Leu Ile
100 105 110
Pro Gly Ile Ser Thr Val Ser Glu Leu Met Leu Gly Met Asp Tyr Gly
115 120 125
Leu Lys Glu Phe Lys Phe Phe Pro Ala Glu Ala Asn Gly Gly Val Lys
130 135 140
Ala Leu Gln Ala Ile Ala Gly Pro Phe Ser Gln Val Arg Phe Cys Pro
145 150 155 160
Thr Gly Gly Ile Ser Pro Ala Asn Tyr Arg Asp Tyr Leu Ala Leu Lys
165 170 175
Ser Val Leu Cys Ile Gly Gly Ser Trp Leu Val Pro Ala Asp Ala Leu
180 185 190
Glu Ala Gly Asp Tyr Asp Arg Ile Thr Lys Leu Ala Arg Glu Ala Val
195 200 205
Glu Gly Ala Lys Leu
210
<210> 4
<211> 642
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(642)
<223> eda
<400> 4
atgaaaaact ggaaaacaag tgcagaatca atcctgacca ccggcccggt tgtaccggtt 60
atcgtggtaa aaaaactgga acacgcggtg ccgatggcaa aagcgttggt tgctggtggg 120
gtgcgcgttc tggaagtgac tctgcgtacc gagtgtgcag ttgacgctat ccgtgctatc 180
gccaaagaag tgcctgaagc gattgtgggt gccggtacgg tgctgaatcc acagcagctg 240
gcagaagtca ctgaagcggg tgcacagttc gcaattagcc cgggtctgac cgagccgctg 300
ctgaaagctg ctaccgaagg gactattcct ctgattccgg ggatcagcac tgtttccgaa 360
ctgatgctgg gtatggacta cggtttgaaa gagttcaaat tcttcccggc tgaagctaac 420
ggcggcgtga aagccctgca ggcgatcgcg ggtccgttct cccaggtccg tttctgcccg 480
acgggtggta tttctccggc taactaccgt gactacctgg cgctgaaaag cgtgctgtgc 540
atcggtggtt cctggctggt tccggcagat gcgctggaag cgggcgatta cgaccgcatt 600
actaagctgg cgcgtgaagc tgtagaaggc gctaagctgt aa 642
<210> 5
<211> 386
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(386)
<223> pheA
<400> 5
Met Thr Ser Glu Asn Pro Leu Leu Ala Leu Arg Glu Lys Ile Ser Ala
1 5 10 15
Leu Asp Glu Lys Leu Leu Ala Leu Leu Ala Glu Arg Arg Glu Leu Ala
20 25 30
Val Glu Val Gly Lys Ala Lys Leu Leu Ser His Arg Pro Val Arg Asp
35 40 45
Ile Asp Arg Glu Arg Asp Leu Leu Glu Arg Leu Ile Thr Leu Gly Lys
50 55 60
Ala His His Leu Asp Ala His Tyr Ile Thr Arg Leu Phe Gln Leu Ile
65 70 75 80
Ile Glu Asp Ser Val Leu Thr Gln Gln Ala Leu Leu Gln Gln His Leu
85 90 95
Asn Lys Ile Asn Pro His Ser Ala Arg Ile Ala Phe Leu Gly Pro Lys
100 105 110
Gly Ser Tyr Ser His Leu Ala Ala Arg Gln Tyr Ala Ala Arg His Phe
115 120 125
Glu Gln Phe Ile Glu Ser Gly Cys Ala Lys Phe Ala Asp Ile Phe Asn
130 135 140
Gln Val Glu Thr Gly Gln Ala Asp Tyr Ala Val Val Pro Ile Glu Asn
145 150 155 160
Thr Ser Ser Gly Ala Ile Asn Asp Val Tyr Asp Leu Leu Gln His Thr
165 170 175
Ser Leu Ser Ile Val Gly Glu Met Thr Leu Thr Ile Asp His Cys Leu
180 185 190
Leu Val Ser Gly Thr Thr Asp Leu Ser Thr Ile Asn Thr Val Tyr Ser
195 200 205
His Pro Gln Pro Phe Gln Gln Cys Ser Lys Phe Leu Asn Arg Tyr Pro
210 215 220
His Trp Lys Ile Glu Tyr Thr Glu Ser Thr Ser Ala Ala Met Glu Lys
225 230 235 240
Val Ala Gln Ala Lys Ser Pro His Val Ala Ala Leu Gly Ser Glu Ala
245 250 255
Gly Gly Thr Leu Tyr Gly Leu Gln Val Leu Glu Arg Ile Glu Ala Asn
260 265 270
Gln Arg Gln Asn Phe Thr Arg Phe Val Val Leu Ala Arg Lys Ala Ile
275 280 285
Asn Val Ser Asp Gln Val Pro Ala Lys Thr Thr Leu Leu Met Ala Thr
290 295 300
Gly Gln Gln Ala Gly Ala Leu Val Glu Ala Leu Leu Val Leu Arg Asn
305 310 315 320
His Asn Leu Ile Met Thr Arg Leu Glu Ser Arg Pro Ile His Gly Asn
325 330 335
Pro Trp Glu Glu Met Phe Tyr Leu Asp Ile Gln Ala Asn Leu Glu Ser
340 345 350
Ala Glu Met Gln Lys Ala Leu Lys Glu Leu Gly Glu Ile Thr Arg Ser
355 360 365
Met Lys Val Leu Gly Cys Tyr Pro Ser Glu Asn Val Val Pro Val Asp
370 375 380
Pro Thr
385
<210> 6
<211> 1161
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(1161)
<223> pheA
<400> 6
atgacatcgg aaaacccgtt actggcgctg cgagagaaaa tcagcgcgct ggatgaaaaa 60
ttattagcgt tactggcaga acggcgcgaa ctggccgtcg aggtgggaaa agccaaactg 120
ctctcgcatc gcccggtacg tgatattgat cgtgaacgcg atttgctgga aagattaatt 180
acgctcggta aagcgcacca tctggacgcc cattacatta ctcgcctgtt ccagctcatc 240
attgaagatt ccgtattaac tcagcaggct ttgctccaac aacatctcaa taaaattaat 300
ccgcactcag cacgcatcgc ttttctcggc cccaaaggtt cttattccca tcttgcggcg 360
cgccagtatg ctgcccgtca ctttgagcaa ttcattgaaa gtggctgcgc caaatttgcc 420
gatattttta atcaggtgga aaccggccag gccgactatg ccgtcgtacc gattgaaaat 480
accagctccg gtgccataaa cgacgtttac gatctgctgc aacataccag cttgtcgatt 540
gttggcgaga tgacgttaac tatcgaccat tgtttgttgg tctccggcac tactgattta 600
tccaccatca atacggtcta cagccatccg cagccattcc agcaatgcag caaattcctt 660
aatcgttatc cgcactggaa gattgaatat accgaaagta cgtctgcggc aatggaaaag 720
gttgcacagg caaaatcacc gcatgttgct gcgttgggaa gcgaagctgg cggcactttg 780
tacggtttgc aggtactgga gcgtattgaa gcaaatcagc gacaaaactt cacccgattt 840
gtggtgttgg cgcgtaaagc cattaacgtg tctgatcagg ttccggcgaa aaccacgttg 900
ttaatggcga ccgggcaaca agccggtgcg ctggttgaag cgttgctggt actgcgcaac 960
cacaatctga ttatgacccg tctggaatca cgcccgattc acggtaatcc atgggaagag 1020
atgttctatc tggatattca ggccaatctt gaatcagcgg aaatgcaaaa agcattgaaa 1080
gagttagggg aaatcacccg ttcaatgaag gtattgggct gttacccaag tgagaacgta 1140
gtgcctgttg atccaacctg a 1161
<210> 7
<211> 108
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(108)
<223> trpR
<400> 7
Met Ala Gln Gln Ser Pro Tyr Ser Ala Ala Met Ala Glu Gln Arg His
1 5 10 15
Gln Glu Trp Leu Arg Phe Val Asp Leu Leu Lys Asn Ala Tyr Gln Asn
20 25 30
Asp Leu His Leu Pro Leu Leu Asn Leu Met Leu Thr Pro Asp Glu Arg
35 40 45
Glu Ala Leu Gly Thr Arg Val Arg Ile Val Glu Glu Leu Leu Arg Gly
50 55 60
Glu Met Ser Gln Arg Glu Leu Lys Asn Glu Leu Gly Ala Gly Ile Ala
65 70 75 80
Thr Ile Thr Arg Gly Ser Asn Ser Leu Lys Ala Ala Pro Val Glu Leu
85 90 95
Arg Gln Trp Leu Glu Glu Val Leu Leu Lys Ser Asp
100 105
<210> 8
<211> 327
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(327)
<223> trpR
<400> 8
atggcccaac aatcacccta ttcagcagcg atggcagaac agcgtcacca ggagtggtta 60
cgttttgtcg acctgcttaa gaatgcctac caaaacgatc tccatttacc gttgttaaac 120
ctgatgctga cgccagatga gcgcgaagcg ttggggactc gcgtgcgtat tgtcgaagag 180
ctgttgcgcg gcgaaatgag ccagcgtgag ttaaaaaatg aactcggcgc aggcatcgcg 240
acgattacgc gtggatctaa cagcctgaaa gccgcgcccg tcgagctgcg ccagtggctg 300
gaagaggtgt tgctgaaaag cgattga 327
<210> 9
<211> 414
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(414)
<223> mtr
<400> 9
Met Ala Thr Leu Thr Thr Thr Gln Thr Ser Pro Ser Leu Leu Gly Gly
1 5 10 15
Val Val Ile Ile Gly Gly Thr Ile Ile Gly Ala Gly Met Phe Ser Leu
20 25 30
Pro Val Val Met Ser Gly Ala Trp Phe Phe Trp Ser Met Ala Ala Leu
35 40 45
Ile Phe Thr Trp Phe Cys Met Leu His Ser Gly Leu Met Ile Leu Glu
50 55 60
Ala Asn Leu Asn Tyr Arg Ile Gly Ser Ser Phe Asp Thr Ile Thr Lys
65 70 75 80
Asp Leu Leu Gly Lys Gly Trp Asn Val Val Asn Gly Ile Ser Ile Ala
85 90 95
Phe Val Leu Tyr Ile Leu Thr Tyr Ala Tyr Ile Ser Ala Ser Gly Ser
100 105 110
Ile Leu His His Thr Phe Ala Glu Met Ser Leu Asn Val Pro Ala Arg
115 120 125
Ala Ala Gly Phe Gly Phe Ala Leu Leu Val Ala Phe Val Val Trp Leu
130 135 140
Ser Thr Lys Ala Val Ser Arg Met Thr Ala Ile Val Leu Gly Ala Lys
145 150 155 160
Val Ile Thr Phe Phe Leu Thr Phe Gly Ser Leu Leu Gly His Val Gln
165 170 175
Pro Ala Thr Leu Phe Asn Val Ala Glu Ser Asn Ala Ser Tyr Ala Pro
180 185 190
Tyr Leu Leu Met Thr Leu Pro Phe Cys Leu Ala Ser Phe Gly Tyr His
195 200 205
Gly Asn Val Pro Ser Leu Met Lys Tyr Tyr Gly Lys Asp Pro Lys Thr
210 215 220
Ile Val Lys Cys Leu Val Tyr Gly Thr Leu Met Ala Leu Ala Leu Tyr
225 230 235 240
Thr Ile Trp Leu Leu Ala Thr Met Gly Asn Ile Pro Arg Pro Glu Phe
245 250 255
Ile Gly Ile Ala Glu Lys Gly Gly Asn Ile Asp Val Leu Val Gln Ala
260 265 270
Leu Ser Gly Val Leu Asn Ser Arg Ser Leu Asp Leu Leu Leu Val Val
275 280 285
Phe Ser Asn Phe Ala Val Ala Ser Ser Phe Leu Gly Val Thr Leu Gly
290 295 300
Leu Phe Asp Tyr Leu Ala Asp Leu Phe Gly Phe Asp Asp Ser Ala Val
305 310 315 320
Gly Arg Leu Lys Thr Ala Leu Leu Thr Phe Ala Pro Pro Val Val Gly
325 330 335
Gly Leu Leu Phe Pro Asn Gly Phe Leu Tyr Ala Ile Gly Tyr Ala Gly
340 345 350
Leu Ala Ala Thr Ile Trp Ala Ala Ile Val Pro Ala Leu Leu Ala Arg
355 360 365
Ala Ser Arg Lys Arg Phe Gly Ser Pro Lys Phe Arg Val Trp Gly Gly
370 375 380
Lys Pro Met Ile Ala Leu Ile Leu Val Phe Gly Val Gly Asn Ala Leu
385 390 395 400
Val His Ile Leu Ser Ser Phe Asn Leu Leu Pro Val Tyr Gln
405 410
<210> 10
<211> 1245
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(1245)
<223> mtr
<400> 10
atggcaacac taaccaccac ccaaacgtca ccgtcgctgc ttggcggcgt ggtgattatc 60
ggcggcacca ttattggcgc agggatgttt tctctgccag tggtcatgtc cggggcgtgg 120
tttttctggt caatggcggc gctgatcttt acctggttct gtatgctgca ttccggcttg 180
atgattctgg aagctaacct gaattacaga atcggttcga gttttgacac catcaccaaa 240
gatttgctgg gcaaaggctg gaacgtggtc aacggcattt ccattgcctt tgtgctctat 300
atcctgacct atgcctatat ttctgccagt ggttcgattc tgcatcacac cttcgcagag 360
atgtcactaa acgtcccggc acgggcggcg ggttttggtt ttgcattgct ggtagcgttt 420
gtggtgtggt tgagcactaa agccgtcagt cgcatgacag cgattgtgct gggggcgaaa 480
gtcattacct tcttcctcac ctttggtagc ctgctggggc atgtgcagcc tgcgacattg 540
ttcaacgtcg ccgaaagcaa tgcgtcttat gcaccgtatc tgttgatgac cctgccgttc 600
tgtctggcat cgtttggtta tcacggtaac gtgccaagcc tgatgaagta ttacggcaaa 660
gatccgaaaa ccatcgtgaa atgtctggtg tacggtacgc tgatggcgct ggcgctgtat 720
accatctggt tgctggcgac gatgggtaac atcccgcgtc cggagtttat cggtattgca 780
gagaagggcg gtaatattga tgtgctggta caggcgttaa gcggcgtact gaacagccgt 840
agtctggatc tgctgctggt cgtgttctca aactttgcgg tagcgagttc gttcctcggc 900
gtaacgctgg gtttgtttga ctatctggca gatctgtttg gtttcgacga ctcggctgtg 960
ggccgcttga aaacggcatt gctgaccttt gccccgccag ttgtgggggg gctgttgttc 1020
ccgaacggat tcctgtacgc cattggttat gctggtttag cggctaccat ctgggcggca 1080
attgttccgg cgctgttagc ccgtgcatcg cgtaaacgct ttggcagccc gaaattccgc 1140
gtctggggtg gcaagccgat gattgcgctg attctggtgt ttggcgtcgg caacgcactg 1200
gtgcatattt tatcgagctt taatttactg ccggtgtatc agtaa 1245
<210> 11
<211> 471
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(471)
<223> tnaA
<400> 11
Met Glu Asn Phe Lys His Leu Pro Glu Pro Phe Arg Ile Arg Val Ile
1 5 10 15
Glu Pro Val Lys Arg Thr Thr Arg Ala Tyr Arg Glu Glu Ala Ile Ile
20 25 30
Lys Ser Gly Met Asn Pro Phe Leu Leu Asp Ser Glu Asp Val Phe Ile
35 40 45
Asp Leu Leu Thr Asp Ser Gly Thr Gly Ala Val Thr Gln Ser Met Gln
50 55 60
Ala Ala Met Met Arg Gly Asp Glu Ala Tyr Ser Gly Ser Arg Ser Tyr
65 70 75 80
Tyr Ala Leu Ala Glu Ser Val Lys Asn Ile Phe Gly Tyr Gln Tyr Thr
85 90 95
Ile Pro Thr His Gln Gly Arg Gly Ala Glu Gln Ile Tyr Ile Pro Val
100 105 110
Leu Ile Lys Lys Arg Glu Gln Glu Lys Gly Leu Asp Arg Ser Lys Met
115 120 125
Val Ala Phe Ser Asn Tyr Phe Phe Asp Thr Thr Gln Gly His Ser Gln
130 135 140
Ile Asn Gly Cys Thr Val Arg Asn Val Tyr Ile Lys Glu Ala Phe Asp
145 150 155 160
Thr Gly Val Arg Tyr Asp Phe Lys Gly Asn Phe Asp Leu Glu Gly Leu
165 170 175
Glu Arg Gly Ile Glu Glu Val Gly Pro Asn Asn Val Pro Tyr Ile Val
180 185 190
Ala Thr Ile Thr Ser Asn Ser Ala Gly Gly Gln Pro Val Ser Leu Ala
195 200 205
Asn Leu Lys Ala Met Tyr Ser Ile Ala Lys Lys Tyr Asp Ile Pro Val
210 215 220
Val Met Asp Ser Ala Arg Phe Ala Glu Asn Ala Tyr Phe Ile Lys Gln
225 230 235 240
Arg Glu Ala Glu Tyr Lys Asp Trp Thr Ile Glu Gln Ile Thr Arg Glu
245 250 255
Thr Tyr Lys Tyr Ala Asp Met Leu Ala Met Ser Ala Lys Lys Asp Ala
260 265 270
Met Val Pro Met Gly Gly Leu Leu Cys Met Lys Asp Asp Ser Phe Phe
275 280 285
Asp Val Tyr Thr Glu Cys Arg Thr Leu Cys Val Val Gln Glu Gly Phe
290 295 300
Pro Thr Tyr Gly Gly Leu Glu Gly Gly Ala Met Glu Arg Leu Ala Val
305 310 315 320
Gly Leu Tyr Asp Gly Met Asn Leu Asp Trp Leu Ala Tyr Arg Ile Ala
325 330 335
Gln Val Gln Tyr Leu Val Asp Gly Leu Glu Glu Ile Gly Val Val Cys
340 345 350
Gln Gln Ala Gly Gly His Ala Ala Phe Val Asp Ala Gly Lys Leu Leu
355 360 365
Pro His Ile Pro Ala Asp Gln Phe Pro Ala Gln Ala Leu Ala Cys Glu
370 375 380
Leu Tyr Lys Val Ala Gly Ile Arg Ala Val Glu Ile Gly Ser Phe Leu
385 390 395 400
Leu Gly Arg Asp Pro Lys Thr Gly Lys Gln Leu Pro Cys Pro Ala Glu
405 410 415
Leu Leu Arg Leu Thr Ile Pro Arg Ala Thr Tyr Thr Gln Thr His Met
420 425 430
Asp Phe Ile Ile Glu Ala Phe Lys His Val Lys Glu Asn Ala Ala Asn
435 440 445
Ile Lys Gly Leu Thr Phe Thr Tyr Glu Pro Lys Val Leu Arg His Phe
450 455 460
Thr Ala Lys Leu Lys Glu Val
465 470
<210> 12
<211> 1416
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(1416)
<223> tnaA
<400> 12
atggaaaact ttaaacatct ccctgaaccg ttccgcattc gtgttattga gccagtaaaa 60
cgtaccactc gcgcttatcg tgaagaggca attattaaat ccggtatgaa cccgttcctg 120
ctggatagcg aagatgtttt tatcgattta ctgaccgaca gcggcaccgg ggcggtgacg 180
cagagcatgc aggctgcgat gatgcgcggc gacgaagcct acagcggcag tcgtagctac 240
tatgcgttag ccgagtcagt gaaaaatatc tttggttatc aatacaccat tccgactcac 300
cagggccgtg gcgcagagca aatctatatt ccggtactga ttaaaaaacg cgagcaggaa 360
aaaggcctgg atcgcagcaa aatggtggcg ttctctaact atttctttga taccacgcag 420
ggccatagcc agatcaacgg ctgtaccgtg cgtaacgtct atatcaaaga agccttcgat 480
acgggcgtgc gttacgactt taaaggcaac tttgaccttg agggattaga acgcggtatt 540
gaagaagttg gtccgaataa cgtgccgtat atcgttgcaa ccatcaccag taactctgca 600
ggtggtcagc cggtttcact ggcaaactta aaagcgatgt acagcatcgc gaagaaatac 660
gatattccgg tggtaatgga ctccgcgcgc tttgctgaaa acgcctattt catcaagcag 720
cgtgaagcag aatacaaaga ctggaccatc gagcagatca cccgcgaaac ctacaaatat 780
gccgatatgc tggcgatgtc cgccaagaaa gatgcgatgg tgccgatggg cggcctgctg 840
tgcatgaaag acgacagctt ctttgatgtg tacaccgagt gcagaaccct ttgcgtggtg 900
caggaaggct tcccgacata tggcggcctg gaaggcggcg cgatggagcg tctggcggta 960
ggtctgtatg acggcatgaa tctcgactgg ctggcttatc gtatcgcgca ggtacagtat 1020
ctggtcgatg gtctggaaga gattggcgtt gtctgccagc aggcgggcgg tcacgcggca 1080
ttcgttgatg ccggtaaact gttgccgcat atcccggcag accagttccc ggcacaggcg 1140
ctggcctgcg agctgtataa agtcgccggt atccgtgcgg tagaaattgg ctctttcctg 1200
ttaggccgcg atccgaaaac cggtaaacaa ctgccatgcc cggctgaact gctgcgttta 1260
accattccgc gcgcaacata tactcaaaca catatggact tcattattga agcctttaaa 1320
catgtgaaag agaacgcggc gaatattaaa ggattaacct ttacgtacga accgaaagta 1380
ttgcgtcact tcaccgcaaa acttaaagaa gtttaa 1416
<210> 13
<211> 415
<212> PRT
<213> Escherichia coli
<220>
<221> PEPTIDE
<222> (1)..(415)
<223> tnaB
<400> 13
Met Thr Asp Gln Ala Glu Lys Lys His Ser Ala Phe Trp Gly Val Met
1 5 10 15
Val Ile Ala Gly Thr Val Ile Gly Gly Gly Met Phe Ala Leu Pro Val
20 25 30
Asp Leu Ala Gly Ala Trp Phe Phe Trp Gly Ala Phe Ile Leu Ile Ile
35 40 45
Ala Trp Phe Ser Met Leu His Ser Gly Leu Leu Leu Leu Glu Ala Asn
50 55 60
Leu Asn Tyr Pro Val Gly Ser Ser Phe Asn Thr Ile Thr Lys Asp Leu
65 70 75 80
Ile Gly Asn Thr Trp Asn Ile Ile Ser Gly Ile Thr Val Ala Phe Val
85 90 95
Leu Tyr Ile Leu Thr Tyr Ala Tyr Ile Ser Ala Asn Gly Ala Ile Ile
100 105 110
Ser Glu Thr Ile Ser Met Asn Leu Gly Tyr His Ala Asn Pro Arg Ile
115 120 125
Val Gly Ile Cys Thr Ala Ile Phe Val Ala Ser Val Leu Trp Ile Ser
130 135 140
Ser Leu Ala Ala Ser Arg Ile Thr Ser Leu Phe Leu Gly Leu Lys Ile
145 150 155 160
Ile Ser Phe Val Ile Val Phe Gly Ser Phe Phe Phe His Val Asp Tyr
165 170 175
Ser Ile Leu Arg Asp Ala Thr Ser Thr Thr Ala Gly Thr Ser Tyr Phe
180 185 190
Pro Tyr Ile Phe Met Ala Leu Pro Val Cys Leu Ala Ser Phe Gly Phe
195 200 205
His Gly Asn Ile Pro Ser Leu Ile Ile Cys Tyr Gly Lys Arg Lys Asp
210 215 220
Lys Leu Ile Lys Ser Val Val Phe Gly Ser Leu Leu Ala Leu Val Ile
225 230 235 240
Tyr Leu Phe Trp Leu Tyr Cys Thr Met Gly Asn Ile Pro Arg Glu Ser
245 250 255
Phe Lys Ala Ile Ile Ser Ser Gly Gly Asn Val Asp Ser Leu Val Lys
260 265 270
Ser Phe Leu Gly Thr Lys Gln His Gly Ile Ile Glu Phe Cys Leu Leu
275 280 285
Val Phe Ser Asn Leu Ala Val Ala Ser Ser Phe Phe Gly Val Thr Leu
290 295 300
Gly Leu Phe Asp Tyr Leu Ala Asp Leu Phe Lys Ile Asp Asn Ser His
305 310 315 320
Gly Gly Arg Phe Lys Thr Val Leu Leu Thr Phe Leu Pro Pro Ala Leu
325 330 335
Leu Tyr Leu Ile Phe Pro Asn Gly Phe Ile Tyr Gly Ile Gly Gly Ala
340 345 350
Gly Leu Cys Ala Thr Ile Trp Ala Val Ile Ile Pro Ala Val Leu Ala
355 360 365
Ile Lys Ala Arg Lys Lys Phe Pro Asn Gln Met Phe Thr Val Trp Gly
370 375 380
Gly Asn Leu Ile Pro Ala Ile Val Ile Leu Phe Gly Ile Thr Val Ile
385 390 395 400
Leu Cys Trp Phe Gly Asn Val Phe Asn Val Leu Pro Lys Phe Gly
405 410 415
<210> 14
<211> 1248
<212> DNA
<213> Escherichia coli
<220>
<221> gene
<222> (1)..(1248)
<223> tnaB
<400> 14
atgactgatc aagctgaaaa aaagcactct gcattttggg gtgttatggt tatagcaggt 60
acagtaattg gtggaggtat gtttgcttta cctgttgatc ttgccggtgc ctggtttttc 120
tggggtgcct ttatccttat cattgcctgg ttttcaatgc ttcattccgg gttattgtta 180
ttagaagcaa atttaaatta tcctgtcggc tccagtttta acaccatcac caaagattta 240
atcggtaaca cctggaacat tatcagcggt attaccgttg ccttcgttct ctatatcctc 300
acttatgcct atatctctgc taatggtgcg atcattagtg aaacgatatc aatgaatttg 360
ggctatcacg ctaatccacg tattgtcggg atctgcacag ccattttcgt tgccagcgta 420
ttgtggataa gttcgttagc cgccagtcgc attacctcat tgttcctcgg gctgaagatt 480
atctcctttg tgatcgtgtt tggttctttc ttcttccatg tcgattactc catcctgcgc 540
gatgccacca gcaccactgc gggaacgtct tacttcccgt atatctttat ggctttgccg 600
gtgtgtctgg cgtcatttgg tttccacggc aatattccca gcctgattat ttgctatgga 660
aaacgcaaag ataagttaat caaaagcgtg gtatttggtt cgctgctggc gctggtgatt 720
tatctcttct ggctctattg cacgatgggg aatattccgc gcgaaagctt taaggcgata 780
atctcctcag gcggcaacgt tgattcgctg gtgaaatcgt tcctcggcac caaacagcac 840
ggcattatcg agttttgcct gctggtgttc tctaacttag ctgttgccag ttcgttcttt 900
ggtgtcacgc tggggttgtt cgattatctg gcggacctgt ttaagattga taactcccac 960
ggcgggcgtt tcaaaaccgt gctgttaacc ttcctgccac ctgcgttgtt gtatctgatc 1020
ttcccgaacg gctttattta cgggatcggc ggtgccgggc tgtgcgccac catctgggcg 1080
gtcattattc ccgcagtgct tgcaatcaaa gctcgcaaga agtttcccaa tcagatgttc 1140
acggtctggg gcggcaatct tattccggcg attgtcattc tctttggtat aaccgtgatt 1200
ttgtgctggt tcggcaacgt ctttaacgtg ttacctaaat ttggctaa 1248
<210> 15
<211> 11155
<212> DNA
<213> Artificial Sequence
<220>
<223> pCL1920_Ptrc_trpO
<400> 15
cccgtcttac tgtcgggaat tcgcgttggc cgattcatta atgcagctgg cacgacaggt 60
ttcccgactg gaaagcgggc agtgagcgca acgcaattaa tgtgagttag ctcactcatt 120
aggcacccca ggctttacac tttatgcttc cggctcgtat gttgtgtgga attgtgagcg 180
gataacaatt tcacacagga aacagctatg accatgatta cgccaagctt gctgttgaca 240
attaatcatc cggctcgtat aatgtgtgga attgtgagcg gataacaatt tcacacagga 300
aacagctgca gatgcaaaca caaaaaccga ctctcgaact gctaacctgc gaaggcgctt 360
atcgcgacaa tcccaccgcg ctttttcacc agttgtgtgg ggatcgtccg gcaacgctgc 420
tgctggaatc cgcagatatc gacagcaaag atgatttaaa aagcctgctg ctggtagaca 480
gtgcgctgcg cattacagct ttaggtgaca ctgtcacaat ccaggcactt tccggcaacg 540
gcgaagccct cctggcacta ctggataacg ccctgcctgc gggtgtggaa agtgaacaat 600
caccaaactg ccgtgtgctg cgcttccccc ctgtcagtcc actgctggat gaagacgccc 660
gcttatgctc cctttcggtt tttgacgctt tccgtttatt gcagaatctg ttgaatgtac 720
cgaaggaaga acgagaagcc atgttcttcg gcggcctgtt ctcttatgac cttgtggcgg 780
gatttgaaga tttaccgcaa ctgtcagcgg aaaataactg ccctgatttc tgtttttatc 840
tcgctgaaac gctgatggtg attgaccatc agaaaaaaag cacccgtatt caggccagcc 900
tgtttgctcc gaatgaagaa gaaaaacaac gtctcactgc tcgcctgaac gaactacgtc 960
agcaactgac cgaagccgcg ccgccgctgc cagtggtttc cgtgccgcat atgcgttgtg 1020
aatgtaatca gagcgatgaa gagttcggtg gcgtagtgcg tttgttgcaa aaagcgattc 1080
gcgctggaga aattttccag gtggtgccat ctcgccgttt ctctctgccc tgcccgtcac 1140
cgctggcggc ctattacgtg ctgaaaaaga gtaatcccag cccgtacatg ttttttatgc 1200
aggataatga tttcacccta tttggcgcgt cgccggaaag ctcgctcaag tatgatgcca 1260
ccagccgcca gattgagatc tacccgattg ccggaacacg cccacgcggt cgtcgcgccg 1320
atggttcact ggacagagat ctcgacagcc gtattgaact ggaaatgcgt accgatcata 1380
aagagctgtc tgaacatctg atgctggttg atctcgcccg taatgatctg gcacgcattt 1440
gcacccccgg cagccgctac gtcgccgatc tcaccaaagt tgaccgttat tcctatgtga 1500
tgcacctcgt ctctcgcgta gtcggcgaac tgcgtcacga tcttgacgcc ctgcacgctt 1560
atcgcgcctg tatgaatatg gggacgttaa gcggtgcgcc gaaagtacgc gctatgcagt 1620
taattgccga ggcggaaggt cgtcgccgcg gcagctacgg cggcgcggta ggttatttca 1680
ccgcgcatgg cgatctcgac acctgcattg tgatccgctc ggcgctggtg gaaaacggta 1740
tcgccaccgt gcaagcgggt gctggtgtag tccttgattc tgttccgcag tcggaagccg 1800
acgaaacccg taacaaagcc cgcgctgtac tgcgcgctat tgccaccgcg catcatgcac 1860
aggagacttt ctgatggctg acattctgct gctcgataat atcgactctt ttacgtacaa 1920
cctggcagat cagttgcgca gcaatgggca taacgtggtg atttaccgca accatattcc 1980
ggcgcaaacc ttaattgaac gcctggcgac catgagcaat ccggtgctga tgctttctcc 2040
tggccccggt gtgccgagcg aagccggttg tatgccggaa ctcctcaccc gcttgcgtgg 2100
caagctgccc attattggca tttgcctcgg acatcaggcg attgtcgaag cttacggggg 2160
ctatgtcggt caggcgggcg aaattctcca cggtaaagcc tccagcattg aacatgacgg 2220
tcaggcgatg tttgccggat taacaaaccc gctgccggtg gcgcgttatc actcgctggt 2280
tggcagtaac attccggccg gtttaaccat caacgcccat tttaatggca tggtgatggc 2340
agtacgtcac gatgcggatc gcgtttgtgg attccagttc catccggaat ccattctcac 2400
cacccagggc gctcgcctgc tggaacaaac gctggcctgg gcgcagcaga aactagagcc 2460
agccaacacg ctgcaaccga ttctggaaaa actgtatcag gcgcagacgc ttagccaaca 2520
agaaagccac cagctgtttt cagcggtggt gcgtggcgag ctgaagccgg aacaactggc 2580
ggcggcgctg gtgagcatga aaattcgcgg tgagcacccg aacgagatcg ccggggcagc 2640
aaccgcgcta ctggaaaacg cagcgccgtt cccgcgcccg gattatctgt ttgctgatat 2700
cgtcggtact ggcggtgacg gcagcaacag tatcaatatt tctaccgcca gtgcgtttgt 2760
cgccgcggcc tgtgggctga aagtggcgaa acacggcaac cgtagcgtct ccagtaaatc 2820
tggttcgtcc gatctgctgg cggcgttcgg tattaatctt gatatgaacg ccgataaatc 2880
gcgccaggcg ctggatgagt taggtgtatg tttcctcttt gcgccgaagt atcacaccgg 2940
attccgccac gcgatgccgg ttcgccagca actgaaaacc cgcaccctgt tcaatgtgct 3000
ggggccattg attaacccgg cgcatccgcc gctggcgtta attggtgttt atagtccgga 3060
actggtgctg ccgattgccg aaaccttgcg cgtgctgggg tatcaacgcg cggcggtggt 3120
gcacagcggc gggatggatg aagtttcatt acacgcgccg acaatcgttg ccgaactgca 3180
tgacggcgaa attaaaagct atcagctcac cgcagaagac tttggcctga caccctacca 3240
ccaggagcaa ctggcaggcg gaacaccgga agaaaaccgt gacattttaa cacgtttgtt 3300
acaaggtaaa ggcgacgccg cccatgaagc agccgtcgct gcgaacgtcg ccatgttaat 3360
gcgcctgcat ggccatgaag atctgcaagc caatgcgcaa accgttcttg aggtactgcg 3420
cagtggttcc gcttacgaca gagtcaccgc actggcggca cgagggtaaa tgatgcaaac 3480
cgttttagcg aaaatcgtcg cagacaaggc gatttgggta gaagcccgca aacagcagca 3540
accgctggcc agttttcaga atgaggttca gccgagcacg cgacattttt atgatgcgct 3600
acagggtgcg cgcacggcgt ttattctgga gtgcaagaaa gcgtcgccgt caaaaggcgt 3660
gatccgtgat gatttcgatc cagcacgcat tgccgccatt tataaacatt acgcttcggc 3720
aatttcggtg ctgactgatg agaaatattt tcaggggagc tttaatttcc tccccatcgt 3780
cagccaaatc gccccgcagc cgattttatg taaagacttc attatcgacc cttaccagat 3840
ctatctggcg cgctattacc aggccgatgc ctgcttatta atgctttcag tactggatga 3900
cgaccaatat cgccagcttg ccgccgtcgc tcacagtctg gagatggggg tgctgaccga 3960
agtcagtaat gaagaggaac aggagcgcgc cattgcattg ggagcaaagg tcgttggcat 4020
caacaaccgc gatctgcgtg atttgtcgat tgatctcaac cgtacccgcg agcttgcgcc 4080
gaaactgggg cacaacgtga cggtaatcag cgaatccggc atcaatactt acgctcaggt 4140
gcgcgagtta agccacttcg ctaacggttt tctgattggt tcggcgttga tggcccatga 4200
cgatttgcac gccgccgtgc gccgggtgtt gctgggtgag aataaagtat gtggcctgac 4260
gcgtgggcaa gatgctaaag cagcttatga cgcgggcgcg atttacggtg ggttgatttt 4320
tgttgcgaca tcaccgcgtt gcgtcaacgt tgaacaggcg caggaagtga tggctgcggc 4380
accgttgcag tatgttggcg tgttccgcaa tcacgatatt gccgatgtgg tggacaaagc 4440
taaggtgtta tcgctggcgg cagtgcaact gcatggtaat gaagaacagc tgtatatcga 4500
tacgctgcgt gaagctctgc cagcacatgt tgccatctgg aaagcattaa gcgtcggtga 4560
aaccctgccc gcccgcgagt ttcagcacgt tgataaatat gttttagaca acggccaggg 4620
tggaagcggg caacgttttg actggtcact attaaatggt caatcgcttg gcaacgttct 4680
gctggcgggg ggcttaggcg cagataactg cgtggaagcg gcacaaaccg gctgcgccgg 4740
acttgatttt aattctgctg tagagtcgca accgggcatc aaagacgcac gtcttttggc 4800
ctcggttttc cagacgctgc gcgcatatta aggaaaggaa caatgacaac attacttaac 4860
ccctattttg gtgagtttgg cggcatgtac gtgccacaaa tcctgatgcc tgctctgcgc 4920
cagctggaag aagcttttgt cagtgcgcaa aaagatcctg aatttcaggc tcagttcaac 4980
gacctgctga aaaactatgc cgggcgtcca accgcgctga ccaaatgcca gaacattaca 5040
gccgggacga acaccacgct gtatctcaag cgtgaagatt tgctgcacgg cggcgcgcat 5100
aaaactaacc aggtgctggg gcaggcgttg ctggcgaagc ggatgggtaa aaccgaaatc 5160
atcgccgaaa ccggtgccgg tcagcatggc gtggcgtcgg cccttgccag cgccctgctc 5220
ggcctgaaat gccgtattta tatgggtgcc aaagacgttg aacgccagtc gcctaacgtt 5280
tttcgtatgc gcttaatggg tgcggaagtg atcccggtgc atagcggttc cgcgacgctg 5340
aaagatgcct gtaacgaggc gctgcgcgac tggtccggta gttacgaaac cgcgcactat 5400
atgctgggca ccgcagctgg cccgcatcct tatccgacca ttgtgcgtga gtttcagcgg 5460
atgattggcg aagaaaccaa agcgcagatt ctggaaagag aaggtcgcct gccggatgcc 5520
gttatcgcct gtgttggcgg cggttcgaat gccatcggca tgtttgctga tttcatcaat 5580
gaaaccaacg tcggcctgat tggtgtggag ccaggtggtc acggtatcga aactggcgag 5640
cacggcgcac cgctaaaaca tggtcgcgtg ggtatctatt tcggtatgaa agcgccgatg 5700
atgcaaaccg aagacgggca gattgaagaa tcttactcca tctccgccgg actggatttc 5760
ccgtctgtcg gcccacaaca cgcgtatctt aacagcactg gacgcgctga ttacgtgtct 5820
attaccgatg atgaagccct tgaagccttc aaaacgctgt gcctgcacga agggatcatc 5880
ccggcgctgg aatcctccca cgccctggcc catgcgttga aaatgatgcg cgaaaacccg 5940
gataaagagc agctactggt ggttaacctt tccggtcgcg gcgataaaga catcttcacc 6000
gttcacgata ttttgaaagc acgaggggaa atctgatgga acgctacgaa tctctgtttg 6060
cccagttgaa ggagcgcaaa gaaggcgcat tcgttccttt cgtcacgctc ggtgatccgg 6120
gcattgagca gtcattgaaa attatcgata cgctaattga agccggtgct gacgcgctgg 6180
agttaggtat ccccttctcc gacccactgg cggatggccc gacgattcaa aacgccactc 6240
tgcgcgcctt tgcggcaggt gtgactccgg cacaatgttt tgaaatgctg gcactgattc 6300
gccagaaaca cccgaccatt cccattggcc tgttgatgta tgccaatctg gtgtttaaca 6360
aaggcattga tgagttttat gcccagtgcg aaaaagtcgg cgtcgattcg gtgctggttg 6420
ccgatgtgcc agttgaagag tccgcgccct tccgccaggc cgcgttgcgt cataatgtcg 6480
cacctatctt catctgcccg ccaaatgccg atgacgacct gctgcgccag atagcctctt 6540
acggtcgtgg ttacacctat ttgctgtcac gagcaggcgt gaccggcgca gaaaaccgcg 6600
ccgcgttacc cctcaatcat ctggttgcga agctgaaaga gtacaacgct gcacctccat 6660
tgcagggatt tggtatttcc gccccggatc aggtaaaagc agcgattgat gcaggagctg 6720
cgggcgcgat ttctggttcg gccattgtta aaatcatcga gcaacatatt aatgagccag 6780
agaaaatgct ggcggcactg aaagtttttg tacaaccgat gaaagcggcg acgcgcagtt 6840
aactgcaggt cgactctaga ggatccccgg gtaccgagct cgaattcact ggccgtcgtt 6900
ttacaacgtc gtgactggga aaaccctggc gttacccaac ttaatcgcct tgcagcacat 6960
ccccctttcg ccagctggcg taatagcgaa gaggcccgca ccgatcgccc ttcccaacag 7020
ttgcgcagcc tgaatggcga atggcgcctg atgcggtatt ttctccttac gcatctgtgc 7080
ggtatttcac accgcatatg gtgcactctc agtacaatct gctctgatgc cgcatagtta 7140
agccagcccc gacacccgcc aacacccgct gacgagctta gtaaagccct cgctagattt 7200
taatgcggat gttgcgatta cttcgccaac tattgcgata acaagaaaaa gccagccttt 7260
catgatatat ctcccaattt gtgtagggct tattatgcac gcttaaaaat aataaaagca 7320
gacttgacct gatagtttgg ctgtgagcaa ttatgtgctt agtgcatcta acgcttgagt 7380
taagccgcgc cgcgaagcgg cgtcggcttg aacgaattgt tagacattat ttgccgacta 7440
ccttggtgat ctcgcctttc acgtagtgga caaattcttc caactgatct gcgcgcgagg 7500
ccaagcgatc ttcttcttgt ccaagataag cctgtctagc ttcaagtatg acgggctgat 7560
actgggccgg caggcgctcc attgcccagt cggcagcgac atccttcggc gcgattttgc 7620
cggttactgc gctgtaccaa atgcgggaca acgtaagcac tacatttcgc tcatcgccag 7680
cccagtcggg cggcgagttc catagcgtta aggtttcatt tagcgcctca aatagatcct 7740
gttcaggaac cggatcaaag agttcctccg ccgctggacc taccaaggca acgctatgtt 7800
ctcttgcttt tgtcagcaag atagccagat caatgtcgat cgtggctggc tcgaagatac 7860
ctgcaagaat gtcattgcgc tgccattctc caaattgcag ttcgcgctta gctggataac 7920
gccacggaat gatgtcgtcg tgcacaacaa tggtgacttc tacagcgcgg agaatctcgc 7980
tctctccagg ggaagccgaa gtttccaaaa ggtcgttgat caaagctcgc cgcgttgttt 8040
catcaagcct tacggtcacc gtaaccagca aatcaatatc actgtgtggc ttcaggccgc 8100
catccactgc ggagccgtac aaatgtacgg ccagcaacgt cggttcgaga tggcgctcga 8160
tgacgccaac tacctctgat agttgagtcg atacttcggc gatcaccgct tccctcatga 8220
tgtttaactt tgttttaggg cgactgccct gctgcgtaac atcgttgctg ctccataaca 8280
tcaaacatcg acccacggcg taacgcgctt gctgcttgga tgcccgaggc atagactgta 8340
ccccaaaaaa acagtcataa caagccatga aaaccgccac tgcgccgtta ccaccgctgc 8400
gttcggtcaa ggttctggac cagttgcgtg agcgcatacg ctacttgcat tacagcttac 8460
gaaccgaaca ggcttatgtc cactgggttc gtgccttcat ccgtttccac ggtgtgcgtc 8520
acccggcaac cttgggcagc agcgaagtcg aggcatttct gtcctggctg gcgaacgagc 8580
gcaaggtttc ggtctccacg catcgtcagg cattggcggc cttgctgttc ttctacggca 8640
aggtgctgtg cacggatctg ccctggcttc aggagatcgg aagacctcgg ccgtcgcggc 8700
gcttgccggt ggtgctgacc ccggatgaag tggttcgcat cctcggtttt ctggaaggcg 8760
agcatcgttt gttcgcccag cttctgtatg gaacgggcat gcggatcagt gagggtttgc 8820
aactgcgggt caaggatctg gatttcgatc acggcacgat catcgtgcgg gagggcaagg 8880
gctccaagga tcgggccttg atgttacccg agagcttggc acccagcctg cgcgagcagg 8940
ggaattaatt cccacgggtt ttgctgcccg caaacgggct gttctggtgt tgctagtttg 9000
ttatcagaat cgcagatccg gcttcagccg gtttgccggc tgaaagcgct atttcttcca 9060
gaattgccat gattttttcc ccacgggagg cgtcactggc tcccgtgttg tcggcagctt 9120
tgattcgata agcagcatcg cctgtttcag gctgtctatg tgtgactgtt gagctgtaac 9180
aagttgtctc aggtgttcaa tttcatgttc tagttgcttt gttttactgg tttcacctgt 9240
tctattaggt gttacatgct gttcatctgt tacattgtcg atctgttcat ggtgaacagc 9300
tttgaatgca ccaaaaactc gtaaaagctc tgatgtatct atctttttta caccgttttc 9360
atctgtgcat atggacagtt ttccctttga tatgtaacgg tgaacagttg ttctactttt 9420
gtttgttagt cttgatgctt cactgataga tacaagagcc ataagaacct cagatccttc 9480
cgtatttagc cagtatgttc tctagtgtgg ttcgttgttt ttgcgtgagc catgagaacg 9540
aaccattgag atcatactta ctttgcatgt cactcaaaaa ttttgcctca aaactggtga 9600
gctgaatttt tgcagttaaa gcatcgtgta gtgtttttct tagtccgtta tgtaggtagg 9660
aatctgatgt aatggttgtt ggtattttgt caccattcat ttttatctgg ttgttctcaa 9720
gttcggttac gagatccatt tgtctatcta gttcaacttg gaaaatcaac gtatcagtcg 9780
ggcggcctcg cttatcaacc accaatttca tattgctgta agtgtttaaa tctttactta 9840
ttggtttcaa aacccattgg ttaagccttt taaactcatg gtagttattt tcaagcatta 9900
acatgaactt aaattcatca aggctaatct ctatatttgc cttgtgagtt ttcttttgtg 9960
ttagttcttt taataaccac tcataaatcc tcatagagta tttgttttca aaagacttaa 10020
catgttccag attatatttt atgaattttt ttaactggaa aagataaggc aatatctctt 10080
cactaaaaac taattctaat ttttcgcttg agaacttggc atagtttgtc cactggaaaa 10140
tctcaaagcc tttaaccaaa ggattcctga tttccacagt tctcgtcatc agctctctgg 10200
ttgctttagc taatacacca taagcatttt ccctactgat gttcatcatc tgagcgtatt 10260
ggttataagt gaacgatacc gtccgttctt tccttgtagg gttttcaatc gtggggttga 10320
gtagtgccac acagcataaa attagcttgg tttcatgctc cgttaagtca tagcgactaa 10380
tcgctagttc atttgctttg aaaacaacta attcagacat acatctcaat tggtctaggt 10440
gattttaatc actataccaa ttgagatggg ctagtcaatg ataattacta gtccttttcc 10500
tttgagttgt gggtatctgt aaattctgct agacctttgc tggaaaactt gtaaattctg 10560
ctagaccctc tgtaaattcc gctagacctt tgtgtgtttt ttttgtttat attcaagtgg 10620
ttataattta tagaataaag aaagaataaa aaaagataaa aagaatagat cccagccctg 10680
tgtataactc actactttag tcagttccgc agtattacaa aaggatgtcg caaacgctgt 10740
ttgctcctct acaaaacaga ccttaaaacc ctaaaggctt aagtagcacc ctcgcaagct 10800
cgggcaaatc gctgaatatt ccttttgtct ccgaccatca ggcacctgag tcgctgtctt 10860
tttcgtgaca ttcagttcgc tgcgctcacg gctctggcag tgaatggggg taaatggcac 10920
tacaggcgcc ttttatggat tcatgcaagg aaactaccca taatacaaga aaagcccgtc 10980
acgggcttct cagggcgttt tatggcgggt ctgctatgtg gtgctatctg actttttgct 11040
gttcagcagt tcctgccctc tgattttcca gtctgaccac ttcggattat cccgtgacag 11100
gtcattcaga ctggctaatg cacccagtaa ggcagcggta tcatcaacag gctta 11155
<210> 16
<211> 38
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 1
<400> 16
aggcaacact atgacatcgt gtaggctgga gctgcttc 38
<210> 17
<211> 40
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 2
<400> 17
ggtcgccatt aacaacgtgg catatgaata tcctccttag 40
<210> 18
<211> 19
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 3
<400> 18
tattgagtgt atcgccaac 19
<210> 19
<211> 18
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 4
<400> 19
cgatgtcata gtgttgcc 18
<210> 20
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 5
<400> 20
ccacgttgtt aatggcgacc 20
<210> 21
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 6
<400> 21
ttcattgaac gggtgatttc 20
<210> 22
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 7
<400> 22
tccgcacgtt tatgatatgc tatcgtactc tttagcgagt acaaccgggg gtgtaggctg 60
gagctgcttc 70
<210> 23
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 8
<400> 23
gccacgtctt atcaggccta caaaatcaat cgcttttcag caacacctct catatgaata 60
tcctccttag 70
<210> 24
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 9
<400> 24
gcgccgggcg tatcgacgca 20
<210> 25
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 10
<400> 25
gcatatcata aacgtgcgga 20
<210> 26
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 11
<400> 26
tgtaggcctg ataagacgtg 20
<210> 27
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 12
<400> 27
aaggggcgat cggcgtgttt 20
<210> 28
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 13
<400> 28
atggcaacac taaccaccac ccaaacgtca ccgtcgctgc ttggcggcgt gtgtaggctg 60
gagctgcttc 70
<210> 29
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 14
<400> 29
ttactgatac accggcagta aattaaagct cgataaaata tgcaccagtg catatgaata 60
tcctccttag 70
<210> 30
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 15
<400> 30
gcagccgtta cattggtaac 20
<210> 31
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 16
<400> 31
gtggtggtta gtgttgccat 20
<210> 32
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 17
<400> 32
tactgccggt gtatcagtaa 20
<210> 33
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 18
<400> 33
tcaaaccgtc agcacggctg 20
<210> 34
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 19
<400> 34
atgaaggatt atgtaatgga aaactttaaa catctccctg aaccgttccg gtgtaggctg 60
gagctgcttc 70
<210> 35
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 20
<400> 35
ttagccaaat ttaggtaaca cgttaaagac gttgccgaac cagcacaaaa catatgaata 60
tcctccttag 70
<210> 36
<211> 21
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 21
<400> 36
ttaagcgaaa tcaccgggga a 21
<210> 37
<211> 18
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 22
<400> 37
atgtccgagc actggcgc 18
<210> 38
<211> 28
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 23
<400> 38
cccaagcttg ctgttgacaa ttaatcat 28
<210> 39
<211> 28
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 24
<400> 39
aaaactgcag ctgtttcctg tgtgaaat 28
<210> 40
<211> 31
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 25
<400> 40
aaaactgcag atgcaaacac aaaaaccgac t 31
<210> 41
<211> 32
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 26
<400> 41
aaaactgcag ttaactgcgc gtcgccgctt tc 32
<210> 42
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 27
<400> 42
aaacgcgttg tgaatcatcc tgctctgaca actcaatttc aggagccttt gccgccagct 60
gaagctttac 70
<210> 43
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 28
<400> 43
acagcacgct tttcagcgcc aggtagtcac ggtagttagc cggagaaata tagtggatct 60
gatgggtacc 70
<210> 44
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 29
<400> 44
tgccctatga gctccggtta caggcgtttc agtcataaat cctctgaatg aaacgcgttg 60
tgaatcatcc 70
<210> 45
<211> 70
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 30
<400> 45
atcgcccgct tccagcgcat ctgccggaac cagccaggaa ccaccgatgc acagcacgct 60
tttcagcgcc 70
<210> 46
<211> 21
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 31
<400> 46
catgatcttg cgcagattgt a 21
<210> 47
<211> 21
<212> DNA
<213> Artificial Sequence
<220>
<223> Primer 32
<400> 47
catgatcttg cgcagattgt a 21
Claims (6)
- 내재적 6-포스포글루코네이트 탈수소효소(6-phosphogluconate dehydratase; edd) 및 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제(2-keto-3-deoxy-6-phosphogluconate aldolase; eda)의 활성이 약화 또는 불활성화되는, L-트립토판 생산능을 갖는 에스케리키아속(Escherichia) 미생물.
- 제1항에 있어서, 상기 6-포스포글루코네이트 탈수소효소는 서열번호 1의 아미노산 서열인, L-트립토판 생산능을 갖는 에스케리키아속 미생물.
- 제1항에 있어서, 상기 2-케토-3-데옥시-6-포스포글루코네이트 알돌라아제는 서열번호 3의 아미노산 서열인, L-트립토판 생산능을 갖는 에스케리키아속 미생물.
- 제1항에 있어서, 상기 에스케리키아속 미생물은 대장균(Escherichia coli)인 것인, L-트립토판 생산능을 갖는 에스케리키아속 미생물.
- 제1항에 있어서, 추가로 pheA, trpR, mtr 및 tnaAB 유전자의 전체 또는 일부가 결실된, L-트립토판 생산능을 갖는 에스케리키아속 미생물.
- 제1항 내지 제5항 중 어느 하나의 항의 에스케리키아속 미생물을 배양하는 단계, 및
상기 배양에 따른 배양물 또는 상기 미생물로부터 L-트립토판을 회수하는 단계를 포함하는, L-트립토판의 제조 방법.
Priority Applications (12)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
KR1020140076698A KR101835173B1 (ko) | 2014-06-23 | 2014-06-23 | L-트립토판 생산능을 갖는 에스케리키아속 미생물 및 이를 이용한 l-트립토판의 제조 방법 |
BR112016030467-5A BR112016030467B1 (pt) | 2014-06-23 | 2015-06-23 | Microrganismo do gênero escherichia produtor de l-triptofano e método para produzir l-triptofano usando o mesmo |
ES15811391T ES2918459T3 (es) | 2014-06-23 | 2015-06-23 | Microorganismo del género Escherichia que produce L-triptófano y método para producir L-triptófano mediante el uso del mismo |
MYPI2016704788A MY172369A (en) | 2014-06-23 | 2015-06-23 | Microorganism of the genus escherichia producing l-tryptophan and method for producing l-tryptophan using the same |
RU2016152662A RU2678139C2 (ru) | 2014-06-23 | 2015-06-23 | Микроорганизм рода Escherichia, продуцирующий L-триптофан, и способ продуцирования L-триптофана с использованием данного микроорганизма |
US15/321,717 US10815510B2 (en) | 2014-06-23 | 2015-06-23 | Microorganism of the genus Escherichia producing L-tryptophan and method for producing L-tryptophan using the same |
CN201580045207.3A CN106574239B (zh) | 2014-06-23 | 2015-06-23 | 生产l-色氨酸的埃希氏菌属微生物和使用其生产l-色氨酸的方法 |
PCT/KR2015/006350 WO2015199406A1 (ko) | 2014-06-23 | 2015-06-23 | L-트립토판 생산능을 갖는 에스케리키아속 미생물 및 이를 이용한 l-트립토판의 제조 방법 |
JP2016575078A JP6687549B2 (ja) | 2014-06-23 | 2015-06-23 | L−トリプトファン生産能を有するエシェリキア属微生物及びこれを用いたl−トリプトファンの製造方法 |
DK15811391.0T DK3159401T3 (da) | 2014-06-23 | 2015-06-23 | Mikroorganisme af slægten Escherichia, der producerer L-tryptophan, og fremgangsmåde til fremstilling af L-tryptophan under anvendelse af samme |
EP15811391.0A EP3159401B1 (en) | 2014-06-23 | 2015-06-23 | Microorganism of the genus escherichia producing l-tryptophan and method for producing l-tryptophan using the same |
JP2018207937A JP2019013256A (ja) | 2014-06-23 | 2018-11-05 | L−トリプトファン生産能を有するエシェリキア属微生物及びこれを用いたl−トリプトファンの製造方法 |
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KR101996767B1 (ko) * | 2018-11-29 | 2019-07-04 | 씨제이제일제당 (주) | cAMP 수용 단백질 변이체 및 이를 이용한 L-아미노산 제조방법 |
KR102134418B1 (ko) * | 2019-06-17 | 2020-07-16 | 씨제이제일제당 주식회사 | L-타이로신을 생산하는 미생물 및 이를 이용한 l-타이로신 생산 방법 |
US20230272364A1 (en) * | 2019-07-11 | 2023-08-31 | Bp Corporation North America Inc. | Gluconate dehydratase enzymes and recombinant cells |
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US913A (en) * | 1838-09-12 | Improved machine for removing snags and sawyers from the beds of rivers | ||
US4371614A (en) * | 1980-08-22 | 1983-02-01 | Ajinomoto Co., Inc. | E.Coli bacteria carrying recombinant plasmids and their use in the fermentative production of L-tryptophan |
JP3932945B2 (ja) * | 2002-03-27 | 2007-06-20 | 味の素株式会社 | L−アミノ酸の製造法 |
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US20050191684A1 (en) * | 2004-02-25 | 2005-09-01 | Zimenkov Danila V. | Method for producing L-amino acids |
RU2004105179A (ru) | 2004-02-25 | 2005-08-10 | Закрытое акционерное общество "Научно-исследовательский институт Аджиномото-Генетика" (ЗАО АГРИ) (RU) | 6-фосфоглюконолактоназа из escherichia coli, фрагмент днк, бактерия, принадлежащая к роду escherichia, -продуцент l-аминокислоты, и способ получения l-аминокислоты |
RU2288268C2 (ru) | 2005-01-26 | 2006-11-27 | Закрытое акционерное общество "Научно-исследовательский институт Аджиномото-Генетика" (ЗАО АГРИ) | 6-фосфоглюконолактоназа из escherichia coli, фрагмент днк, бактерия, принадлежащая к роду escherichia - продуцент l-аминокислоты и способ получения l-аминокислоты |
KR100792095B1 (ko) | 2006-12-29 | 2008-01-04 | 씨제이 주식회사 | L-페닐알라닌 생산능을 갖는 대장균 변이주로부터유전자조작된 l-트립토판 생산능을 갖는 재조합 대장균균주 및 이를 이용한 트립토판 제조방법 |
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WO2009101757A1 (ja) * | 2008-02-14 | 2009-08-20 | Panasonic Corporation | コンデンサマイクロホン及びmemsデバイス |
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DK2803720T3 (da) * | 2012-01-10 | 2022-04-04 | Cj Cheiljedang Corp | Mikroorganismer fra escherichia coli med forbedret produktion af l-tryptofan og fremgangsmåde til fremstilling af l-tryptofan dermed |
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JP2017518759A (ja) | 2017-07-13 |
US10815510B2 (en) | 2020-10-27 |
RU2016152662A (ru) | 2018-07-23 |
WO2015199406A1 (ko) | 2015-12-30 |
US20170137854A1 (en) | 2017-05-18 |
DK3159401T3 (da) | 2022-07-04 |
CN106574239B (zh) | 2024-05-14 |
CN106574239A (zh) | 2017-04-19 |
ES2918459T3 (es) | 2022-07-15 |
MY172369A (en) | 2019-11-21 |
EP3159401A4 (en) | 2018-01-10 |
EP3159401A1 (en) | 2017-04-26 |
BR112016030467B1 (pt) | 2023-11-14 |
JP2019013256A (ja) | 2019-01-31 |
EP3159401B1 (en) | 2022-05-18 |
RU2016152662A3 (ko) | 2018-07-23 |
BR112016030467A2 (ko) | 2018-01-16 |
JP6687549B2 (ja) | 2020-04-22 |
KR101835173B1 (ko) | 2018-03-07 |
RU2678139C2 (ru) | 2019-01-23 |
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