KR20000068633A - 당전이 활성을 갖는 단백질을 암호화하는 유전자 - Google Patents
당전이 활성을 갖는 단백질을 암호화하는 유전자 Download PDFInfo
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- KR20000068633A KR20000068633A KR1019997002547A KR19997002547A KR20000068633A KR 20000068633 A KR20000068633 A KR 20000068633A KR 1019997002547 A KR1019997002547 A KR 1019997002547A KR 19997002547 A KR19997002547 A KR 19997002547A KR 20000068633 A KR20000068633 A KR 20000068633A
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Abstract
본 발명은 서열 7 내지 10 또는 12중 어느 하나에 기재된 아미노산 서열을 가지며 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자와 상기 아미노산 서열에 대하여 변형된 아미노산 서열을 갖는 동시에 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자 및 당해 유전자를 사용하는 상기 단백질의 제조방법을 제공한다. 이러한 유전자는 식물의 색의 인공적 개량 등을 위해 사용할 수 있다.
Description
화훼 산업은 신규한 각종 품종을 개발하는 데에 노력하고 있다. 신규한 품종의 육성을 위한 효과적인 방법의 하나로서 꽃의 색을 변색시키는 경우가 있으며 고전적인 육종 방법을 사용하여 대부분의 상업적 품종에 대해서 광범위한 색을 생성하는 데에 성공하고 있다. 그러나, 이 방법으로는 품종마다 유전자 풀이 제한되어 있으므로 단일한 품종이 광범위한 종류의 착색 품종을 갖는 경우는 희귀하다.
꽃의 색은 주로 두가지 타입의 색소, 즉 플라보노이드 및 카로티노이드에 근거하며 플라보노이드는 황색으로부터 적색 내지 청색의 범위에 기여하고 카로티노이드는 오렌지 또는 황색의 색조에 기여한다. 꽃의 색으로 주된 기여를 하는 플라보노이드 분자는 시아니딘, 델피니딘, 페튜니딘, 페오니딘, 말비딘 및 펠라르고니딘의 배당체인 안토시아닌이며 상이한 안토시안이 현저한 꽃의 색 변화를 일으킨다. 또한 꽃의 색은 무색의 플라보노이드의 보조 발색, 금속 착체 형성, 글루코실화, 아실화, 메틸화 및 액포의 pH에 따라 영향받는다(참조: Forkmann, Plant Breeding, l06, 1, l991).
페닐알라닌으로부터 시작되는 안토시아닌의 생합성 경로는 잘 이해되어 있으며(예: P1ant Cell, 7, l071-1083, 1995) 생합성에 관계되는 유전자는 대부분 클로닝되어 있다. 예를 들면, 자소(Perilla)의 안토시아닌인 말로닐시소닌(3-0-(6-0-(p-쿠마로일)-β-D-글루코실)-5-0-(6-0-말로닐-β-D-글루코실)-시아니딘)의 생합성에 관계한다고 간주되는 유전자 중에서 이의 동족체가 현재까지 보고되어 있지 않은 것은 플라보노이드-3'-하이드록실라제, UDP-글루코스:안토시아닌(플라보노이드) 5-0-글루코실 트란스페라제(이하, 5GT), 말로닐기 트란스페라제 유전자뿐이다.
이중에서 플라보노이드-3'-하이드록실라제는 시토크롬 P450 유전자군에 속하는 것으로 공지되어 있으며(참조: Plant Ce11, 7, 1071-1083, 1995) 시토크롬 P450 유전자는 서로 구조적인 상동성을 나타내는 것으로 고찰된다.
일반적으로 플라보노이드 분자의 3위치의 하이드록실 그룹은 글루코스에 의해 개질되지만 글루코실화를 위시해서 배당체에 의한 개질은 안토시아닌의 안정성과 용해성을 증대시킨다고 생각된다(참조: The Flavonoids, Chapman & Hall, 1994).
이러한 반응을 촉매하는 UDP-글루코스:안토시아니딘 또는 플라보노이드-3-글루코실트란스페라제(이하, 3GT)를 암호화하는 유전자는 옥수수, 대맥, 금어초 및 용담 등의 많은 식물로부터 수득되고 있으며 아미노산 서열은 서로 상당한 상동성을 나타낸다. 예를 들면, 단자엽 식물인 옥수수와 쌍자엽 식물인 용담의 3GT의 아미노산 서열의 상동성은 32%, 단자엽 식물인 옥수수와 대맥의 3GT의 아미노산 서열의 상동성은 73%, 쌍자엽 식물인 용담과 가지의 3GT의 아미노산 서열에서는 46%이다.
또한, 페튜니아의 UDP-람노스:안토시아니딘-3-글루코사이드람노실트란스페라제(3RT)를 암호화하는 유전자도 클로닝되어 있다.
그러나, 많은 식물의 플라보노이드의 5위치의 하이드록실 그룹이 글루코실화되는 것에 관계없이 이러한 반응을 촉매하는 효소(5GT)의 유전자는 아직도 수득되지 않고 있다.
또한, 페튜니아나 및 스톡의 안토시아닌의 5위치로 배당체를 전이하는 반응을 측정한 예는 있지만(참조: Planta l60, 341-347, 1984, Planta, 168, 586-591, 1986) 이들 보고는 꽃잎의 조추출액이나 부분 정제한 것을 사용하여 효소학적 성질을 조사하는 데에 머물고 있으며 이러한 효소를 순수한 형태로 정제한 예는 없다. 또한, 일반적으로 글리코실트란스페라제는 생화학적으로 불안정하며 효소의 정제가 곤란하다.
플라보노이드 분자에 배당체가 부가함에 따른 색조의 변화는 거의 없지만 색조에 큰 영향을 주는 방향족 아실기는 안토시아닌 내의 글루코스 분자나 람노스 분자로 전이하므로 배당체 전이 반응의 조절은 안토시아닌의 생합성을 제어하며 궁극적으로는 꽃의 색을 제어하는 데에 있어서 중요하다. 또한 글리코실트란스페라제 효소 유전자의 발현을 조절하여 꽃의 색을 바꾼 예로서 페튜니아의 3RT에 따른 반응을 형질 전환된 페튜니아에서 제어하여 꽃의 색을 개질한 예가 있다.
외래 유전자로 형질 전환할 수 있는 식물로서는 예를 들면, 장미, 국화, 카네이션, 가베라, 페튜니아, 토레니아, 터키 도라지, 카란코에, 튜우립, 글라디올라스 등이 공지되어 있다.
발명의 개시
따라서, 본 발명자 등은 플라보노이드의 5위치로 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자를 얻는 것을 과제로 하여 본 발명을 완성하였다.
예를 들면, 국화의 안토시아닌 및 장미 및 카네이션의 안토시아닌의 일부는 5위치의 하이드록실 그룹이 글루코실화되어 있지 않다. 본 발명으로 얻어진 5GT 유전자를 이들의 식물에 도입함으로써 안토시아닌의 구조를 변형시킬 수 있다.
또한, 국제공개공보 WO 96/25500에 기재되어 있는 아실기 트란스페라제 유전자를 사용하여 플라보노이드를 아실화함으로써 꽃색을 변화시키거나 플라보노이드를 안정화시킬 수 있지만 아실기는 직접 플라보노이드와 결합하는 것이 아니고 당을 통하여 결합하므로 아실기 트란스페라제 유전자를 도입한 것만으로는 꽃의 색 변화가 충분하지 않거나 안정화되지 않는 경우도 있다.
그러나, 아실기 트란스페라제 유전자와 동시에 5GT 유전자를 도입함으로써 플라보노이드의 5위치에 슈가를 결합시킨 다음, 이를 아실화할 수 있으며 이로써 안토시아닌의 구조가 바뀌고 꽃의 색이 푸르게 되는 것도 예상할 수 있다.
또한, 안토시아닌의 5위치가 글루코실화되어 있는 식물의 5GT 유전자의 발현을 안티센스법이나 코-서프레션(co-suppression)법 등으로 억제하면 5 위치로의 글루코스 잔기의 전이를 억제할 수 있으며 그 결과, 꽃의 색을 변화시킬 수 있다. 예를 들면, 용담이나 터키 도라지에서 5GT 활성을 억제하면 꽃의 색은 붉게 될 수 있다.
본 발명자는 유전자 재조합기술을 사용하여 자소, 토레니아, 버베나 및 페튜니아로부터 5GT의 cDNA를 단리하여 구조 유전자의 염기 서열을 결정하였다. 즉, 본 발명자는 이들 식물에서 안토시아닌을 발현하는 조직에 존재하는 5GT를 암호화하는 DNA 서열을 제공하였다. 또한, 이러한 효소는 안토시아닌 색소의 5위치로 배당체를 전이시키므로 꽃의 색 변화에 이용할 수 있으며 안토시아닌의 안정성을 증가시킬 수 있다.
발명의 실시의 형태
본 발명의 효소를 암호화하는 DNA를 얻는 데는 예를 들면, 디퍼런셜 디스플레이(Differential display) 법을 사용할 수 있다. 예를 들면, 자소(Peril1a frutescens)에 있어서는 안토시아닌을 축적하는 품종[예: 자훈(紫熏)]과 안토시아닌을 축적하지 않은 품종[예: 청훈(靑熏)]이 있으며 안토시아닌을 축적하는 품종에는 존재하지만 안토시아닌을 축적하지 않은 품종에는 존재하지 않는 DNA를 클로닝하면 본 발명의 효소를 암호화하는 DNA가 얻어질 가능성이 있다.
보다 구체적으로는 자훈의 잎 및 청훈의 잎으로부터 RNA를 추출하고 통상적인 방법에 따라 cDNA를 합성하고 이것을 전기영동으로 분리하여 자훈 유래의 cDNA 라이브러리 내에 존재하고 청훈 유래의 cDNA 라이브러리 내에는 존재하지 않는 cDNA를 단리한다. 다음에 이와 같이 얻어지는 cDNA를 프로브로서 사용하여 자훈 유래의 cDNA 라이브러리를 스크리닝하여 본 발명의 효소를 암호화하는 DNA를 얻는다.
상기와 같이 하여 본 발명의 효소를 암호화하는 cDNA가 얻어지면 이러한 cDNA 또는 이의 단편을 프로브로서 사용하여 다른 식물로부터 cDNA 라이브러리를 스크리닝함으로써 당해 식물 유래의 본 발명의 효소를 암호화하는 DNA를 얻을 수 있다.
본 발명에서 상기한 스크리닝의 예로서 디퍼런셜 디스플레이에 의해 자소 유래의 본 발명의 효소를 암호화하는 DNA를 클로닝하고 (실시예 1) 다음에 이와 같이 얻어진 DNA를 프로브로서 버베나(Verbena hybrida)로부터 cDNA를 스크리닝함으로써 본 발명의 효소를 암호화하는 DNA를 버베나로부터 얻으며(실시예 2) 다시 동일하게 하여 토레니아로부터 본 발명의 효소를 암호화하는 DNA를 얻는다(실시예 3).
그리고, 이들 DNA가 본 발명의 효소 활성을 갖는 단백질을 발현하는 것을 확인한다.
또한, 본 발명의 효소를 암호화하는 DNA를 페튜니아로부터 얻는다(실시예 4).
본 발명의 DNA로서는 예를 들면, 서열 7 내지 10 또는 12중의 어느 하나에 기재된 아미노산 서열을 암호화하는 것을 들 수 있다. 그러나, 복수개의 아미노산의 부가, 결실 및/또는 하나 이상의 다른 아미노산에 의한 치환에 따라 개질된 아미노산 서열을 갖는 단백질도 원래의 단백질과 동일한 효소 활성을 유지하는 것으로 공지되어 있다. 따라서 본 발명은 서열 7 내지 l0 또는 l2중의 어느 하나에 기재된 아미노산 서열에 대하여 1개 또는 복수개의 아미노산의 부가, 결실 및/또는 다른 아미노산으로 치환된 개질된 아미노산 서열을 가지며 또한, 플라보노이드의 5위치로 배당체를 전이하는 활성을 유지하는 단백질을 암호화하는 유전자도 본 발명에 속한다.
본 발명은 또한, 2 내지 5×SSC, 예를 들면, 5×SSC, 50℃의 조건하에서 서열 1 내지 4 또는 6중의 어느 하나에 기재된 염기 서열 또는 여기에 기재된 아미노산 서열을 암호화하는 염기 서열 또는 이들 부분, 예를 들면, 컨센서스 영역의 6개 이상의 아미노산을 암호화하는 부분과 하이브리드화하는 동시에 플라보노이드의 5위치에 배당체를 전이시키는 활성을 갖는 단백질을 암호화하는 유전자에 관한 것이다. 또한, 최적인 하이브리드화 온도는 염기 서열이나 이의 길이에 따라 상이하고 염기 서열이 짧아짐에 따라 하이브리드화 온도는 낮게 하는 것이 바람직하며 예를 들면, 아미노산 6개를 암호화하는 염기 서열(l8 염기)의 경우에 50℃ 이하의 온도가 바람직하다.
이러한 하이브리드화에 의해 선택되는 유전자로서는 천연 유래의 것, 예를 들면, 식물 유래의 것, 예를 들면, 버베나, 토레니아 유래의 유전자를 들 수 있지만 다른 식물, 예를 들면, 페튜니아, 장미, 카네이션, 히아신스 등에서 유래하는 유전자도 포함한다. 또한, 하이브리드화에 의해 선택되는 유전자는 cDNA 또는 게놈 DNA일 수 있다.
본 발명은 또한, 서열 7 내지 10 또는 12중의 어느 하나에 기재된 아미노산 서열에 대하여 30% 이상, 바람직하게는 50% 이상, 예를 들면, 60% 또는 70% 이상, 경우에 따라서는 90% 이상의 상동성을 갖는 아미노산 서열을 갖는 동시에 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자에 관한 것이다. 즉, 실시예에 기재된 바와 같이 본 발명의 효소를 암호화하는 DNA는 다른 글리코실트란스페라제 유전자와 비교하여 20 내지 30%의 상동성을 나타낸다. 따라서, 본 발명은 서열 7 내지 10 또는 12에 기재된 아미노산 서열과 30% 이상의 상동성을 나타내며 또한 글리코실트란스페라제 활성을 갖는 단백질을 암호화하는 유전자를 포함한다.
또한, 실시예 1 내지 4의 결과의 비교에서 명백한 바와 같이 본 발명의 효소의 아미노산 서열은 품종에 따라 상이하며 품종간의 상동성은 50% 이상(참조: 실시예 3 및 4), 예를 들면, 60 내지 70%(참조: 실시예 2)이고 또한 동일 품종 유래의 효소 아미노산 서열의 상동성은 90% 이상(참조: 실시예 1)이다. 따라서 본 발명은 서열 7 내지 10 또는 12에 기재된 아미노산 서열에 대하여 50% 이상, 예를 들면, 60 내지 70% 이상, 경우에 따라서는 90% 이상의 상동성을 갖는 아미노산 서열을 가지며 또한 본 발명의 글리코실트란스페라제 효소 활성을 유지하는 단백질을 암호화하는 유전자도 본 발명의 범위이다.
생물 유래의 염기 서열을 갖는 DNA는 실시예에 구체적으로 기재한 바와 같이 예를 들면, cDNA 라이브러리의 스크리닝에 의해 얻어진다.
또한, 개질된 아미노산 서열을 갖는 효소를 암호화하는 DNA는 생물 유래의 염기 서열을 갖는 DNA를 기초하여 통상적인 부위-특이적 돌연변이 및 PCR법을 사용하여 합성할 수 있다. 예를 들면, 변이시키고자 하는 부위를 함유하는 DNA 단편을 상기에 의해 얻어지는 cDNA 또는 게놈 DNA의 제한 효소 분해에 의해 얻을 수 있으며 이것을 주형으로 하여 원하는 변이를 포함하는 프라이머를 사용하여 부위 특이적 돌연변이 또는 PCR 법을 실시하고 원하는 변이를 포함하는 DNA 단편을 얻고 이를 표적 효소의 다른 부분을 암호화하는 DNA에 연결시킨다.
또는, 단축된 아미노산 서열을 갖는 효소를 암호화하는 DNA를 수득하기 위해 예를 들면, 표적 아미노산 서열보다 긴 아미노산 서열, 예를 들면, 전체 길이의 아미노산 서열을 암호화하는 DNA를 원하는 제한 효소로 절단하여 얻어진 DNA 단편이 수득된 아미노산 서열의 전체를 암호화하지 않는 경우에는 합성 DNA를 연결함으로써 결실 부분을 보충해야만 한다.
또한, 이러한 클론을 이.콜리(E.coli) 또는 효모에서 유전자 발현 시스템을 사용하여 발현시키고 효소 활성을 측정함으로써 얻어진 유전자가 글리코실트란스페라제를 암호화하는 것을 확인하고 플라보노이드의 5위치에 배당체를 전이시키는 글리코실트란스페라제 유전자의 해독 영역을 명백히 함으로써 본 발명에 따른 글리코실트란스페라제를 암호화하는 유전자가 얻어지며 또한, 당해 유전자를 발현시킴으로써 유전자 산물인, 플라보노이드의 5위치에 배당체를 전이시키는 표적 글리코실트란스페라제 단백질을 얻을 수 있다.
또한, 서열 7 내지 10 또는 12중의 어느 하나에 기재된 아미노산 서열에 대한 항체를 사용하여도 상기한 단백질을 얻을 수 있다.
따라서 본 발명은 또한 상기한 DNA를 함유하는 재조합 벡터, 특히 발현 벡터 및 당해 벡터에 의해 형질 전환되는 숙주에 관한 것이다. 숙주로서는 원핵 생물 또는 진핵 생물을 사용할 수 있다. 원핵 생물로서는 세균, 예를 들면 에스케리치아(Escherichia)속에 속하는 세균, 예를 들면, 에스케리치아 콜리(Escherichia coli), 바실러스(Bacillus)속에 속하는 세균, 예를 들면, 바실러스 서브틸리스(Bacillus subtilis) 등의 통상적인 숙주를 사용할 수 있다.
진핵성 숙주로서는 하등 진핵생물과 같은 진핵성 미생물, 예를 들면, 진균류인 효모 또는 곰팡이를 사용할 수 있다. 효모로서는 예를 들면, 사카로마이세스(Saccharomyces)속 미생물, 예를 들면, 사카로마이세스 세레비지에 (Saccharomyces cerevisiae) 등을 들 수 있으며 또한 곰팡이로서는 아스퍼질러스 (Aspergillus)속 미생물, 예를 들면, 아스퍼질러스 오리제(Aspergillus oryzae) 및 아스퍼질러스 나이거(Aspergillus niger)등이 있으며 또한 페니실리움 (Penicillium)속 미생물 등을 들 수 있다. 또한, 동물 세포 또는 식물 세포를 사용할 수 있으며 동물 세포로서는 마우스, 햄스터, 원숭이 및 사람 세포계가 사용된다. 또한, 곤충 세포, 예를 들면, 누에의 세포 또는 누에의 성충 자체도 숙주로서 사용된다.
본 발명의 발현 벡터는 이들을 도입해야 할 숙주의 종류에 의존하여 발현 조절 영역, 예를 들면, 프로모터 및 터미네이터, 복제 오리진 등을 함유한다. 세균 발현 벡터의 프로모터로서는 통상적인 프로모터, 예를 들면, trc 프로모터, tac 프로모터 및 1ac 프로모터 등이 사용되며 효모용 프로모터로서는 예를 들면, 글리세로알데히드 트리포스페이트 데하이드로게나제 프로모터 및 PH05 프로모터 등이 사용되며 곰팡이용 프로모터로서는 예를 들면, 아밀라제 및 trp C 등이 사용된다. 또한, 동물세포 숙주용 프로모터로서는 바이러스성 프로모터, 예를 들면, SV 40 얼리 프로모터 및 SV40 레이트 프로모터 등이 사용된다.
발현 벡터의 작제는 제한 효소 및 리가제 등을 사용하여 통상적인 방법에 따라 실시할 수 있다. 또한, 발현 벡터를 사용하는 숙주의 형질 전환도 통상적인 방법에 따라 실시할 수 있다.
상기한 단백질의 제조방법에서는 상기한 발현 벡터로 형질 전환된 숙주를 배양, 재배 또는 사육하고 배양물 등으로부터 통상적인 방법에 따라 예를 들면, 여과, 원심분리, 세포의 파쇄, 겔여과 크로마토그래피, 이온 교환 크로마토그래피 등에 의해 표적 단백질을 회수 및 정제할 수 있다.
또한, 본 명세서에서는 자소, 버베나, 토레니아 및 페튜니아 유래의, 플라보노이드의 5위치에 배당체를 전이시키는 트란스페라제(본 발명에서 단순히 「글리코실트란스페라제」라고 언급됨)에 관해 기재되어 있으며 당해 효소의 정제법을 그대로 또는 일부를 변형하여 다른 식물의 글리코실트란스페라제를 정제하고 당해 효소에 따른 아미노산 서열을 결정함으로써 당해 효소를 암호화하는 유전자를 클로닝할 수 있다. 또한, 본 발명에 따른 자소 유래의 글리코실트란스페라제의 cDNA를 프로브로서 사용함으로써 자소로부터 상이한 글리코실트란스페라제의 cDNA 및 다른 식물로부터 상이한 글리코실트란스페라제의 cDNA를 얻을 수 있다. 따라서, 글리코실트란스페라제의 유전자의 일부 또는 전부를 사용하면 다른 글리코실트란스페라제 유전자를 얻을 수 있다.
또한, 본 명세서에 기재된 바와 같이 자소, 버베나, 토레니아 및 페튜니아 유래의 글리코실트란스페라제를 정제하여 통상적인 방법에 따라 당해 효소에 대한 항체를 얻음으로써 클로닝될 수 있는 이러한 항체와 반응하는 단백질을 만드는 cDNA 또는 염색체 DNA를 클로닝할 수 있다.
따라서, 본 발명은 자소, 버베나, 토레니아 및 페튜니아 유래의 글리코실트란스페라제의 유전자만으로 한정되는 것이 아니며 광범위한 글리코실트란스페라제에 관한 것이다.
또한 본 발명은 글리코실트란스페라제의 유전자를 도입함으로써 색이 조절되는 식물 또는 이의 자손 또는 이들의 조직에 관한 것이며 당해 형태는 절화일 수 있다.
또한, 본 명세서에서는 안토시아닌을 포함한 플라보노이드의 배당체 전이 반응에서 배당체의 공여체로서 UDP-글루코스를 들 수 있다.
본 발명은 플라보노이드의 5위치에 당(glycoside)을 전이하는 활성을 갖는 단백질을 암호화하는 유전자 및 이의 이용방법에 관한 것이다.
하기에 본 발명을 실시예에 기초하여 상세하게 설명한다. 실험 순서는 특별히 기술하지 않는 한, 문헌[참조: Molecular Cloning, Cold Spring Harbor, l989), 신세이부쓰가가쿠짓켄노안나이 제3권(가가쿠도진 1996), 국제공개공보; WO96/25500]에 기재된 방법에 따른다.
실시예 1. 자훈에서 특이적으로 발현되는 유전자의 클로닝
(1) 디퍼런셜 디스플레이
자소(Perilla frutescens)에는 잎에 안토시아닌을 축적하는 품종[예: 자훈(사카타의 혈통)]과 안토시아닌을 축적하지 않는 품종[예: 청훈(사카타의 혈통)]이 있으며 주요한 안토시아닌의 구조는 말로닐시소닌(3-0-(6-0-(p-쿠마로 일)-β-D-글루코실)5-0-(6-0말로닐-β-D-글루코실)-시아니딘]인 것이 보고되어 있다(참조: Agri. Biol. Chem. 53:197-198, 1989).
디퍼런셜 디스플레이는 문헌[Science 257, 967-97l(1992)]에 보고된 방법이며 조직 특이적으로 발현하는 유전자를 얻는 데 사용되고 있다.
상기한 2종류의 자소의 잎으로부터 고온 페놀법(참조: Plant Molecular Biology Manual, Kluwer Academic Publishers l994 pp. D5/1-13)에 의해 전체 RNA를 추출한다. 얻어진 전체 RNA에서 mRNA 분리 키트(Clonetech사)를 사용하여 폴리 A + RNA를 정제한다. 0.9μg의 폴리 A + RNA를 앵커를 부가한 올리고 dT 프라이머(GenHunter사의 H-T11G, H-T11A, H-TllC)를 사용하여 반응 혼합물 33μ1에서 역전사하여 1본쇄 cDNA를 얻는다. 이러한 cDNA를 주형으로 하고 앵커를 부가한 동일한 올리고 dT 프라이머와 합성 프라이머(GenHunter사의 H-APl 내지 8)를 프라이머로 하고 PCR을 실시한다.
PCR의 반응 혼합물의 부피는 20μ1이며 2μ1의 cDNA 용액, 0.2μM의 H-TllG, H-T11A, H-T11C중의 하나의 프라이머, 0.2μM의 H-AP1 내지 8중의 하나의 프라이머, 0.12μM dNTP, 5 또는 10μCi의 [32P] dCTP, l0mM Tris-HC1(pH9.0), 50mM KC1, 0.01% 트리톤 X-100, 1.25mM MgCl2및 1 유니트의 Taq 폴리메라제를 함유하고 있다. 반응 조건은 하기와 같다. 72℃에서 20초 동안 유지한 다음, 94℃에서 30초, 40℃에서 2분, 72℃에서 30초를 l 사이클로 하는 반응을 40 사이클 반복하여 72℃에서 5분 동안 유지한다.
상기와 같이 증폭된 DNA 단편을 DNA 염기 서열화를 위해 사용되는 것과 동일한 폴리아크릴아미드겔 전기영동으로 분리한다. 겔을 건조한 다음, X선 필름에서 노출시킨다. 얻어진 밴드 약 2,600개중에서 2종류의 품종의 자소를 비교하면 자훈에서만 보여지는 밴드는 36개이다. 이들을 건조한 겔로부터 절취하여 l00μ1의 물에 용출시킨다. 용출된 DNA를 에탄올로 침전시켜 20μ1의 물에 용해한다. 이중에서 반량의 DNA를 주형으로 하고 상기에 기재한 PCR 반응을 각각 실시하고 33종류의 밴드에 대한 DNA 단편을 증폭할 수 있다. 이러한 DNA 단편을 사용하여 라이브러리의 스크리닝과 노던 분석을 실시한다.
(2) 노던 분석
상기한 33종류의 DNA 프로브를 사용하여 하기의 방법으로 노던 분석을 실시한다. 자훈과 청훈 유래의 폴리 A + RNA를 1.2% 아가로스를 함유하는 포름아미드 겔로 분리한 다음, 나일론 막에 옮긴다. 이러한 막을 5XSSPE, 5X 덴할트액, 0.5% SDS, 20μg/ml의 변성 연어 정자 DNA 존재하에 65℃에서 하룻밤 동안32P로 표지한 상기 DNA 프로브와 하이브리드화시킨다. 하이브리드화된 막을 1 XSSPE 및 0.1% SDS 용액 속에서 65℃에서 세정하여 오토라디오그래피를 얻는다. 그 결과, 5종류의 프로브만이 자훈에서 특이적으로 발현한다. 이들 클론은 안토시아닌의 생합성에 관계하는 유전자인 것으로 예상된다.
(3) cDNA 라이브러리의 스크리닝
자훈의 잎으로부터 얻은 폴리 A + RNA를 사용하고 컴플리트 래피드 클로닝 시스템 λgtl0(Amersham)를 사용하여 λgtl0을 벡터로 하는 cDNA 라이브러리를 작제한다. 이러한 cDNA 라이브러리를 상기한 5종류의 DNA 단편을 사용하여 스크리닝하고 각각의 단편에 상응하는 cDNA를 얻는다. 이중에서 3R5라는 명칭의 클론은 H-T11A와 H-AP3의 프라이머로부터 유래된 DNA 단편을 사용하여 수득되며, 이미 보고되어 있는 옥수수의 플라보노이드-3-0-글리코실트란스페라제와 아미노산 수준에서 약 26%의 상동성을 나타낸다.
또한, 동일한 프로브를 사용하는 라이브러리의 스크리닝으로 3R4 및 3R6으로 명명된 클론이 얻어지며 이들은 3R5와 대단히 높은 상동성을 나타낸다. 3R4 및 3R6의 전체 염기 서열과 추정된 아미노산 서열을 각각 서열목록의 서열 1과 서열 2에 기재한다. 또한 3R4와 3R6에 암호화된 단백질의 추정 아미노산 서열은 92%의 상동성을 나타낸다.
8R6이라고 명명된 클론은 HT11G와 H-AP8의 프라이머로부터 유래된 DNA 단편을 사용하여 얻어진 것이며 지금까지 보고되어 있는 DNA 염기 서열과는 상당한 상동성을 나타내지 않는다. 이러한 서열을 서열목록의 서열 5에 나타낸다. 8R6은 안토시아닌의 생합성에 관계하는 유전자일 가능성이 강하지만 이의 구조가 지금까지 보고되어 있는 유전자와 상동성이 없으므로 안토시아닌 생합성에 관계하는 신규 유전자인 것으로 예상된다.
자소의 안토시아닌(상기한 말로닐시소닌)의 구조를 고려하면 본 유전자는 말로닐 트란스페라제인 것으로 예상된다. 이를 증명하기 위해 이러한 유전자를 효모나 이 콜리에서 발현시키고 안토시아닌과 말로닐 CoA를 기질로서 반응시켜야만 한다. 이러한 실험은 예를 들면, 국제공개공보 W0 96/25500에 기재되어 있는 방법을 사용하여 실시할 수 있다. 말로닐기 트란스페라제 유전자도 안토시아닌의 구조를 인위적으로 변화시키는 데에 있어서 유용하다.
(4) 효모에 있어서 3R4의 cDNA의 발현
p3R4의 BstX1 절단 부위를 T4 DNA 폴리메라제(Takara Shuzo)를 사용하여 평활화한 다음, 어댑터 내의 BamHI 절단 부위에서 절단 인출하여 얻어지는 약 1.5kb의 DNA 단편과 pYE 22m의 EcoRI 절단 말단을 평활화하고 다시 BamHI로 분해하여 얻어지는 약 8kb의 DNA 단편을 연결하여 얻어지는 플라스미드를 pY3R4로 한다.
또한, pYE 22m을 갖는 이 콜리 균주 JM l09는 에스케리치아 콜리 SBM 335라고 명명하고 FERM BP-5435로서 고교기쥬쓰인세이메이고가쿠고교기쥬쓰겐큐쇼에 기탁되어 있다. pY3R4에서 글리코실트란스페라제를 암호화하는 cDNA는 효모의 구성적인 프로모터의 하나인 글리세로알데히드 트리포스페이트 데하이드로게나제 효소의 프로모터의 하부에 연결되어 있으며 상기한 프로모터로 전사가 조절된다.
pY3R4를 사용하여 효모 사카로마이세스 세레비지에(Saccharomyces cerevi siae) Gl315(Ashikari et al.,Appl. Microbiol. Biotechnol., 30, 515-520, 1989)를 이토 등의 방법(Ito, et a1., J. Bacteriol., 153, 163-168,1983)으로 형질 전환한다. 형질 전환된 효모는 트립토판의 합성능력의 회복에 따라 선택한다. 얻어진 형질 전환된 균주를 10ml의 1% 카자미노산(Difco사)을 함유하는 버크홀더 배지 (Burkholder, Amer. J. Bot. 30, 206-2l0)에서 30℃로 24시간 동안 진탕 배양한다.
아울러서 대조 실험을 위해 트립토판의 합성능력을 자발적으로 회복한 효모도 동일하게 배양한다. 이들을 집균후에 현탁 완충액(100mM 인산염 완충액(pH 8.5), 0.1%(v/v) 2-머캅토에탄올, l0μM APMSF, 100μM UDP-글루코스)에 현탁하고 유리 비드(Glass Beads 425600microns Acid-Wash, sigma사)를 가하여 격렬하게 진탕함으로써 세포를 분쇄한다. 분쇄된 세포를 15,000rpm에서 20분 동안 원심분리한 상등액을 조효소액으로 하고 하기의 효소 활성측정에 사용한다.
(5) 효소활성의 측정
조효소액 20μ1를 함유하는 50μ1 반응 혼합물[l00mM 인산염 완충액(pH 8.5), 670μM 시아니딘-3-글루코사이드, lmM UDP-글루코스]을 30℃에서 l0분 동안 반응시킨 다음, 0.1% TFA를 함유하는 50% 아세트니트릴 용액 50μ1을 첨가하여 반응을 정지시킨다. 15,000rpm에서 5분 동안 원심분리한 상등액을 삼프렙 LCR4(T)-LC 필터(Millipore)를 통해 불순물을 제거한다. 이것을 고속 액체 크로마토그래피 (HPLC)로 분석한다. 분석은 역상 칼럼(Asahipak 0DP50, 4.6mm 직경 x 250mm, Showa Denko)을 사용하고 이동상은 A 용액(0.5% TFA/H2O) 및 B 용액(0.5% TFA 50% CH2CN)으로 구성되고 유속은 0.6ml/min이며 분획물은 B 20%→ B l00%(20분)의 농도구배로 용출시킨 후에 B l00%에서 5분 동안 유지시킨다.
분석에는 반응 용액 20μl를 사용한다. A 520nm, AUFS 0.5(Shimadzu SPD-l0A)와 포토다이오드 어레이 검출기(Shimadzu SPD-M6A)를 사용하여 600-250nm의 흡광도로 검출한다. pY3R4를 발현시킨 효모의 조효소액을 반응시키는 경우, 기질 시아니딘-3-글루코사이드(체류 시간: 17분)뿐만 아니라 14.5분의 체류 시간을 갖는 새로운 피크가 관찰된다. 이것은 대조 실험의 효모의 조효소액를 반응시키는 경우에 관찰되지 않으므로 새로운 피크는 pY3R4로부터 유래하는 단백질의 활성으로 인해 발생한다고 생각된다. 시아니딘-3,5-디글루코사이드와의 공동의 크로마토그래피의 결과, 이러한 피크의 체류 시간은 시아니딘-3,5-디글루코사이드와 일치하며 또한 양쪽의 흡수 스펙트럼도 일치한다. 상기한 관찰로부터 자소의 3R4의 cDNA가 5GT을 암호화한다는 것을 알았다.
실시예 2. 버베나(Verbena hybrida)의 5GT 유전자의 클로닝
(1) cDNA 라이브러리의 작제
버베나 품종 화수국 바이올레트(Suntory)로부터 꽃잎을 모으고 액체 질소내에서 유발로 연마 분쇄한다. 이러한 연마된 분쇄된 조직으로부터 구아니딘티오시아네이트/염화세슘을 사용하는 방법으로 RNA를 추출하고 올리고텍스(Takara Shuzo)를 사용하여 제조자가 권장하는 방법으로 폴리 A + RNA를 얻는다. 구아니딘티오시아네이트/염화세슘을 사용하는 방법은 문헌[참조: Methods in Molecular Biology vol 2, (Humana Press, Inc. 1984) by R.McGookin and Robert J.Slater]에 상세하게 기재되어 있는 방법에 따른다.
수득된 폴리 A + RNA를 주형으로 하고 ZAP-cDNA 합성 키트(Stratagene)를 사용하여 2본쇄 cDNA를 합성한 다음, Uni-ZAP XR 클로닝 키트(Stratagene)를 사용하여 제조자의 권장하는 방법으로 cDNA 라이브러리를 작제한다.
(2) 5GT의 cDNA의 클로닝
상기와 같이 하여 얻어지는 λ파지 라이브러리를 자소의 p3R4의 cDNA를 프로브로 하여 아래와 같이 스크리닝한다. 필터를 하이브리드화 완충액[5X SSC, 30% 포름아미드, 50mM 인산나트륨 완충액(pH 7.0), 3% SDS, 2% 차단제(Boehringer), 0.1% 라우로일사르코신, 80μg/m1 연어 정자 DNA] 속에서 42℃로 1시간 동안 유지한다. DIG 표지된 자소의 5GT cDNA, p3R4의 cDNA 단편을 하이브리드화 용액에 가한 다음, 필터를 16시간 동안 항온처리한다.
세정액(5 X SSC 50℃, 1% SDS)으로 필터를 세정한 다음, 알칼리 포스파타제로 표지된 DIG 특이적인 항체를 사용하는 효소면역 측정법(베링거사)에 따라 5-브로모-4-클로로3인돌릴포스페이트와 니트로 블루 테트라졸륨염의 발색 반응으로 프로브가 하이브리드화된 양성 클론을 검출한다. 검출 방법은 사용설명서에 따른다.
그 결과, 7개의 양성 클론이 얻어진다. 스트라타진사의 권장하는 방법으로 이들 cDNA를 플라스미드 pB1uescript SK로부터 절단 회수한다. 아가로스겔 전기영동으로 cDNA의 길이를 조사한 바, 최장 2.0kb의 삽입이 확인된다.
(3) 염기 서열의 결정
얻어진 클론로부터 플라스미드를 추출하여 서열 분석기 ABI 373A(Perkin-Elmer)를 사용하고 상기 회사가 권장하는 형광 시약을 사용하는 디데옥시 서열 방법으로 cDNA의 3' 및 5' 말단 부근의 염기 서열을 결정한다. 그 결과, 이들 7개의 클론중에 5개의 클론은 서로 동일한 염기서열을 가지고 있으며 cDNA의 길이가 상이한 것으로 생각된다. 이중에서 pSHGT 8의 전체 염기서열을 결정한다. 염기 서열의 결정은 킬로-서열 검출 키트(Takara Shuzo)를 사용하여 일련의 결실 클론을 얻거나 또는 pSHGT 8의 내부 서열에 특이적인 올리고 프라이머를 사용하여 상기와 같이 실시한다.
(4) 염기 서열과 아미노산 서열의 비교
pSHGT 8에 삽입된 cDNA는 2062 bp이고 그 속에 1386 bp(정지 코돈을 함유한다)로 이루어진 오픈 리딩 프레임(ORF)을 포함한다. 이러한 서열을 서열 3에 기재한다. 이러한 ORF의 아미노산 서열은 자소의 p3R4에 암호화된 5GT의 아미노산 서열과 68% 및 p3R6에 암호화되는 것과 64%의 상동성을 나타낸다. 또한, 단자엽 식물 및 쌍자엽 식물의 3GT와 22 내지 25% 및 페튜니아의 3RT와 2l%의 상동성을 나타낸다.
(5) 효모에서의 발현과 효소활성의 측정
pSHGT 8을 BamHI/XhoI로 절단하여 얻어지는 약 2.0kb의 DNA 단편과 pYE 22m을 BamHI/SalI로 절단하여 얻어지는 약 8kb의 DNA 단편을 연결하여 얻어지는 플라스미드를 pYHGT 8로 명명한다. 실시예 1과 동일한 방법으로 효모 균체내에서 pYHGT 8을 발현하고 pSHGT 8에 의해 암호화되는 단백질의 효소 활성에 관해 측정한다. 그 결과, pYHGT 8로 형질전환된 효모의 조효소액을 포함하는 반응 혼합물에서 시아니딘-3,5-디글루코사이드와 체류 시간 및 흡수 스펙트럼이 일치하는 산물을 수득한다. 상기 관찰을 토대로 버베나의 pSHGT 8의 cDNA는 5GT를 암호화한다는 것을 알 수 있다.
실시예 3. 토레니아의 5GT 유전자의 클로닝
(1) cDNA 라이브러리의 작제
토레니아 품종 썸머 웨이브 블루[Suntory]로부터 꽃잎을 모아 액체 질소중에서 유발로 연마 분쇄한다. 이러한 연마 분쇄 조직물로부터 구아니딘티오시아네이트 /염화세슘을 사용하는 방법에 의해 RNA를 추출하고 올리고텍스[Takara Shuzo]를 사용하여 제조자가 권장하는 방법으로써 폴리 A + RNA를 얻는다. 구아니딘티오시아네이트/염화세슘을 사용하는 방법은 문헌[참조: Methods in Molecular Bio1ogy, vo1 2(Humana Press Inc. 1984) by R.McGookin and Robert J, Slater, et al]에 상세하게 기재되어 있는 방법에 따른다.
수득된 폴리 A + RNA를 주형으로 하고 스트라타진사의 ZAP-cDNA 합성 키트를 사용하여 2본쇄 cDNA를 합성한 다음, Uni-ZAP XR 클로닝 키트(Stratagene)를 사용하여 제조자가 권장하는 방법으로 cDNA 라이브러리를 작제한다.
(2) 5GT의 cDNA의 클로닝
상기와 같이 하여 얻어진 λ파지 라이브러리를 자소의 p3R4의 cDNA를 프로브로서 실시예 2와 동일하게 스크리닝한다. 이러한 결과, 8개의 양성 클론이 얻어진다. cDNA를 플라스미드 pBluescript SK 상에서 절단 회수한 다음, 아가로스겔 전기영동으로 cDNA의 길이를 조사한 바, 최장 1.6kb의 삽입이 확인된다.
(3) 염기서열의 결정
얻어진 클론으로부터 플라스미드를 추출하고 실시예 2와 동일하게 하여 5' 및 3'의 양쪽 말단 부근의 염기서열을 결정한다. 그 결과, 이들 클론 중에서 6개는 서로 동일한 염기 서열을 가지며 cDNA의 길이가 다른 것으로 생각된다. 이러한 6 클론 중에서 pSTGT 5의 cDNA의 전체 염기서열을 결정한다.
(4) 염기 서열과 아미노산 서열의 비교
pSTGT 5에 삽입된 cDNA는 167lbp이고 그 속에 1437bp(정지 코돈을 함유한다)로 이루어진 오픈 리딩 프레임(ORF)을 포함한다. 이러한 서열을 서열 4에 나타낸다. 이러한 ORF의 아미노산 서열은 자소의 p3R4에 의해 암호화되는 5GT의 아미노산 서열과 58%, p3R6에 의해 암호화되는 것과 57% 및 버베나의 pSHGT 8에 의해 암호화되는 것과 57%의 상동성을 나타낸다. 또한, 단자엽 식물 및 쌍자엽 식물의 3GT와 19 내지 23% 및 페튜니아의 3RT와 20%의 상동성을 나타낸다.
(5) 효모에서의 5GT 유전자의 발현
pSTGT 5를 SmaI/KpnI로 절단하여 얻어지는 약 1.6kb의 DNA 단편과 pYE 22m의 EcoRI 절단 부위를 평활화한 다음, KpnI로 분해하여 얻어지는 약 8kb의 DNA 단편을 연결하여 얻어지는 플라스미드를 pYTGT 5로 명명한다. 실시예 1과 동일한 방법으로 효모 균체내에 pYTGT 5를 발현하여 pSTGT 5에 암호화되는 단백질의 효소 활성을 측정한다. 그 결과, pYTGT 5로 형질전환된 효모의 조효소액를 포함하는 반응 혼합물에서 시아니딘-3,5-디글루코사이드와 체류시간 및 흡수 스펙트럼 모두 일치하는 생성물이 얻수득된다. 이러한 관찰을 토대로 토레니아의 pSTGT 5의 cDNA는 5GT를 암호화한다는 것을 알았다.
실시예 4. 페튜니아의 5GT 유전자의 클로닝
(1) cDNA 라이브러리의 작제
페튜니아 품종 0ld Glory Blue의 꽃잎으로부터 추출한 RNA를 기초하여 문헌[참조: T. Ho1ton, et al., (Plant Journal, 1993 4:1003-1010)]에 상세하게 기재되어 있는 바와 같이 cDNA 라이브러리를 작제한다.
(2) 5GT의 cDNA의 클로닝
상기와 같이 하여 얻어진 자소, 토레니아, 버베나의 5GT cDNA 혼합물을 프로브로 하여 실시예 2와 동일한 방법으로 스크리닝한다. 이 결과, 얻어진 양성 클론중에서 4개를 플라스미드 pBluescript SK 상에 절단 회수한다. 아가로스 전기영동으로 cDNA의 길이를 조사한 바, 최장 2.0kb의 cDNA가 확인된다.
(3) 염기 서열의 결정
플라스미드를 수득한 클론으로부터 추출한다. 5' 말단 부근의 염기 서열을 실시예 2와 동일한 방법으로 결정한다. 그 결과, 이들 클론중에서 2개의 pSPGT l은 지금까지 얻어진 자소, 토레니아, 버베나의 5GT와 높은 상동성을 보이는 아미노산 서열을 암호화하는 것이 명백해졌다. 따라서 pSPGT l의 전체 염기 서열을 결정한다.
(4) 염기 서열과 아미노산 서열의 비교
pSPGT 1에 삽입된 cDNA는 2015bp이고 이중에서 1407bp(정지 코돈을 함유한다)로 이루어진 오픈 리딩 프레임(ORF)을 포함한다. 이러한 서열을 서열 6에 나타낸다. 이러한 ORF의 아미노산 서열은 자소의 p3R4에 암호화되는 5GT의 아미노산 서열과 57%, p3R6에 암호화되는 아미노산 서열과 54%, 버베나의 pSHGT 8에 암호화되는 것과 55% 및 토레니아의 pTGT5에 암호화되는 것과 51%의 상동성을 나타낸다. 또한 단자엽 식물, 쌍자엽 식물의 3GT와 20 내지 29% 및 페튜니아의 3RT와 20%의 상동성을 나타낸다. 이러한 점으로부터 페튜니아로부터 얻어진 pSPGT1의 cDNA는 5GT를 암호화한다고 생각된다.
상기와 같이 자소, 버베나, 토레니아 및 페튜니아 유래의, 플라보노이드의 5위치에 배당체를 전이하는 효소를 암호화하는 cDNA를 클로닝시키고 이의 염기 서열을 결정한다. 또한, 효모에서 발현되는 이의 단백질의 효소 활성으로 보아 분리된 cDNA가 5GT를 암호화한다는 것이 명백하다. 이러한 cDNA를 적당한 식물 발현 벡터에 도입하여 수득한 발현 작제물을 식물에 도입하고 형질전환된 세포내의 5GT의 활성을 부여하거나 증가시키거나 감소시킴으로써 식물의 꽃 색상 조절에 이용할 수 있게 되었다. 또한, 본 효소를 이용함으로써 식물 내에서 또는 시험관 내에서 안토시안의 구조를 변형시켜 보다 안정적인 안토시안을 합성할 수 있다.
서열
서열: 1
서열의 길이: 1507
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 자소(Perilla frutescens)
조직의 종류: 잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: p3R4
서열
서열: 2
서열의 길이: 1470
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 자소(Perilla frutescens)
조직의 종류: 잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: p3R6
서열
서열: 3
서열의 길이: 2062
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 바베나(Verbena hybrida)
조직의 종류: 꽃잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: pSHGT8
서열
서열: 4
서열의 길이: 1671
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 토레니아
조직의 종류: 꽃잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: pSTGT5
서열
서열: 5
서열의 길이: 1437
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 자소(Perilla frutescens)
조직의 종류: 잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: p8R6
서열
서열: 6
서열의 길이: 2105
서열의 형: 핵산
쇄의 수: 2본쇄
토로포지: 직쇄상
기원
생물명: 페튜니아
조직의 종류: 꽃잎
직접기원
라이브러리명: cDNA 라이브러리
클론명: pSPGT1
서열
〈110〉 Suntory Limited
〈120〉 Gene coding for protein having sugar-transfer activity
〈130〉 5-1998-096135-9
〈150〉 PCT/JP 97/200571
〈151〉 1997-07-25
〈160〉 12
〈170〉 KOPATIN 1.0
〈210〉 1
〈211〉 1507
〈212〉 DNA
〈213〉 Perilla frutescens
〈220〉
〈221〉 CDS
〈222〉 (17)..(1396)
〈400〉 1
gaaaatttcc acaaaa atg gtc cgc cgc cgc gtg ctg cta gca acg ttt 49
Met Val Arg Arg Arg Val Leu Leu Ala Thr Phe
1 5 10
cct gcg caa ggc cac ata aat ccc gcc ctc caa ttc gcc aag aga ctc 97
Pro Ala Gln Gly His Ile Asn Pro Ala Leu Gln Phe Ala Lys Arg Leu
15 20 25
cta aaa gcc ggc act gac gtc aca ttt ttc acg agc gtt tat gca tgg 145
Leu Lys Ala Gly Thr Asp Val Thr Phe Phe Thr Ser Val Tyr Ala Trp
30 35 40
cgc cgc atg gcc aac aca gcc tcc gcc gct gcc gga aac cca ccg ggc 193
Arg Arg Met Ala Asn Thr Ala Ser Ala Ala Ala Gly Asn Pro Pro Gly
45 50 55
ctc gac ttc gtg gcg ttc tcc gac ggc tac gac gac ggg ctg aag ccc 241
Leu Asp Phe Val Ala Phe Ser Asp Gly Tyr Asp Asp Gly Leu Lys Pro
60 65 70 75
tgc ggc gac ggg aag cgc tac atg tcc gag atg aaa gcc cgc ggc tcc 289
Cys Gly Asp Gly Lys Arg Tyr Met Ser Glu Met Lys Ala Arg Gly Ser
80 85 90
gag gcc tta aga aac ctc ctt ctc aac aac cac gac gtc acg ttc gtc 337
Glu Ala Leu Arg Asn Leu Leu Leu Asn Asn His Asp Val Thr Phe Val
95 100 105
gtc tac tcc cac ctc ttt gca tgg gcg gcg gag gtg gcg cgt gag tcc 385
Val Tyr Ser His Leu Phe Ala Trp Ala Ala Glu Val Ala Arg Glu Ser
110 115 120
cag gtc ccg agc gcc ctt ctc tgg gtc gag ccc gcc acc gtg ctg tgc 433
Gln Val Pro Ser Ala Leu Leu Trp Val Glu Pro Ala Thr Val Leu Cys
125 130 135
ata tat tac ttc tac ttc aac ggc tac gca gac gag atc gac gcc ggt 481
Ile Tyr Tyr Phe Tyr Phe Asn Gly Tyr Ala Asp Glu Ile Asp Ala Gly
140 145 150 155
tcc gac gaa att cag ctc cct cgg ctt cca ccc ctg gag cag cgc agt 529
Ser Asp Glu Ile Gln Leu Pro Arg Leu Pro Pro Leu Glu Gln Arg Ser
160 165 170
ctt ccg acc ttt ctg ctg ccg gag aca ccg gag aga ttc cgg ttg atg 577
Leu Pro Thr Phe Leu Leu Pro Glu Thr Pro Glu Arg Phe Arg Leu Met
175 180 185
atg aag cag aag ctg gaa act tta gac ggt gaa gag aag gcg aaa gtg 625
Met Lys Gln Lys Leu Glu Thr Leu Asp Gly Glu Glu Lys Ala Lys Val
190 195 200
ttg gtg aac acg ttt gat gcg ttg gag ccc gat gca ctc acg gct att 673
Leu Val Asn Thr Phe Asp Ala Leu Glu Pro Asp Ala Leu Thr Ala Ile
205 210 215
gat agg tat gag ttg atc ggg atc ggg ccg ttg att ccc tcc gcc ttc 721
Asp Arg Tyr Glu Leu Ile Gly Ile Gly Pro Leu Ile Pro Ser Ala Phe
220 225 230 235
ttg gac ggc gga gat ccc tcc gaa acg tct tac ggc ggc gat ctt ttc 769
Leu Asp Gly Gly Asp Pro Ser Glu Thr Ser Tyr Gly Gly Asp Leu Phe
240 245 250
gaa aaa tcg gag gag aat aac tgc gtg gag tgg ttg gac acg aag ccg 817
Glu Lys Ser Glu Glu Asn Asn Cys Val Glu Trp Leu Asp Thr Lys Pro
255 260 265
aaa tct tcg gtg gtg tat gtg tcg ttt ggg agc gtt ttg agg ttt cca 865
Lys Ser Ser Val Val Tyr Val Ser Phe Gly Ser Val Leu Arg Phe Pro
270 275 280
aag gca caa atg gaa gag att ggg aaa ggg cta tta gcc tgc gga agg 913
Lys Ala Gln Met Glu Glu Ile Gly Lys Gly Leu Leu Ala Cys Gly Arg
285 290 295
ccg ttt tta tgg atg ata cga gaa cag aag aat gac gac ggc gaa gaa 961
Pro Phe Leu Trp Met Ile Arg Glu Gln Lys Asn Asp Asp Gly Glu Glu
300 305 310 315
gaa gaa gaa gag ttg agt tgc att ggg gaa ttg aaa aaa atg ggg aaa 1009
Glu Glu Glu Glu Leu Ser Cys Ile Gly Glu Leu Lys Lys Met Gly Lys
320 325 330
ata gtt tcg tgg tgc tcg cag ttg gag gtt ctg gcg cac cct gcg ttg 1057
Ile Val Ser Trp Cys Ser Gln Leu Glu Val Leu Ala His Pro Ala Leu
335 340 345
gga tgt ttc gtg acg cat tgt ggg tgg aac tcg gct gtg gag agc ttg 1105
Gly Cys Phe Val Thr His Cys Gly Trp Asn Ser Ala Val Glu Ser Leu
350 355 360
agt tgc ggg gtt ccg gtg gtg gcg gtg ccg cag tgg ttt gat cag acg 1153
Ser Cys Gly Val Pro Val Val Ala Val Pro Gln Trp Phe Asp Gln Thr
365 370 375
acg aat gcg aag ctg att gag gat gcg tgg ggg aca ggg gtg aga gtg 1201
Thr Asn Ala Lys Leu Ile Glu Asp Ala Trp Gly Thr Gly Val Arg Val
380 385 390 395
aga atg aat gaa ggg ggt ggg gtt gat gga tct gag ata gag agg tgt 1249
Arg Met Asn Glu Gly Gly Gly Val Asp Gly Ser Glu Ile Glu Arg Cys
400 405 410
gtg gag atg gtg atg gat ggg ggt gag aag agc aaa cta gtg aga gaa 1297
Val Glu Met Val Met Asp Gly Gly Glu Lys Ser Lys Leu Val Arg Glu
415 420 425
aat gcc ata aaa tgg aag act ttg gcc aga gaa gcc atg gga gag gat 1345
Asn Ala Ile Lys Trp Lys Thr Leu Ala Arg Glu Ala Met Gly Glu Asp
430 435 440
gga tct tca ctc aag aat ctc aac gcc ttt ctt cat caa gtt gca cgt 1393
Gly Ser Ser Leu Lys Asn Leu Asn Ala Phe Leu His Gln Val Ala Arg
445 450 455
gct taat acacaaaatg gctttccact tttaatctac tcaaacaccg gttcaaataa 1450
Ala
460
atatcccctt ccacttcttt ctatttcact atcacattta taattttagt aacaaaa 1507
〈210〉 2
〈211〉 1470
〈212〉 DNA
〈213〉 Perilla frutescens
〈220〉
〈221〉 CDS
〈222〉 (28)..(1353)
〈400〉 2
accaaaccaaa acaaaatttc cacaaaa atg gtc cgc cgc cgc gtg ctg cta 48
Met Val Arg Arg Arg Val Leu Leu
1 5
gca acg ttt ccg gcg caa ggc cac ata aat ccc gcc ctc caa ttc gcc 96
Ala Thr Phe Pro Ala Gln Gly His Ile Asn Pro Ala Leu Gln Phe Ala
10 15 20
aag aga ctc cta aaa gcc ggc act gac gtc acg ttt ttc acg agc gtt 144
Lys Arg Leu Leu Lys Ala Gly Thr Asp Val Thr Phe Phe Thr Ser Val
25 30 35 40
tat gca tgg cgc cgc atg gcc aac aca gcc tcc gcc gct gcc gga aac 192
Tyr Ala Trp Arg Arg Met Ala Asn Thr Ala Ser Ala Ala Ala Gly Asn
45 50 55
cca ccg ggc ctc gac ttc gtg gcg ttc tcc gac ggc tac gac gac ggg 240
Pro Pro Gly Leu Asp Phe Val Ala Phe Ser Asp Gly Tyr Asp Asp Gly
60 65 70
ctg aag ccc ggc ggc gac ggg aag cgc tac atg tcc gag atg aaa gcc 288
Leu Lys Pro Gly Gly Asp Gly Lys Arg Tyr Met Ser Glu Met Lys Ala
75 80 85
cgc ggc tcc gag gcc tta aga aac ctc ctt ctc aac aac gac gac gtc 336
Arg Gly Ser Glu Ala Leu Arg Asn Leu Leu Leu Asn Asn Asp Asp Val
90 95 100
act ttc gtc gtc tac tcc cac ctc ttt gca tgg gcg gcg gag gtg gcg 384
Thr Phe Val Val Tyr Ser His Leu Phe Ala Trp Ala Ala Glu Val Ala
105 110 115 120
cgt ttg tcc cac gtc ccg acc gcc ctt ctc tgg gtc gag ccc gcc acc 432
Arg Leu Ser His Val Pro Thr Ala Leu Leu Trp Val Glu Pro Ala Thr
125 130 135
gtg ctg tgc ata tac cac ttc tac ttc aac ggc tac gca gac gag atc 480
Val Leu Cys Ile Tyr His Phe Tyr Phe Asn Gly Tyr Ala Asp Glu Ile
140 145 150
gac gcc ggt tcc aat gaa att cag ctc cct cgg ctt cca tcc ctg gag 528
Asp Ala Gly Ser Asn Glu Ile Gln Leu Pro Arg Leu Pro Ser Leu Glu
155 160 165
cag cgc agt ctt ccg acg ttt ctg ctg cct gcg acg ccg gag aga ttc 576
Gln Arg Ser Leu Pro Thr Phe Leu Leu Pro Ala Thr Pro Glu Arg Phe
170 175 180
cgg ttg atg atg aag gag aag ctg gaa act tta gac ggt gaa gag aag 624
Arg Leu Met Met Lys Glu Lys Leu Glu Thr Leu Asp Gly Glu Glu Lys
185 190 195 200
gcg aaa gta ttg gtg aac acg ttt gat gcg ttg gag ccc gat gca ctc 672
Ala Lys Val Leu Val Asn Thr Phe Asp Ala Leu Glu Pro Asp Ala Leu
205 210 215
acg gct att gat agg tat gag ttg atc ggg atc ggg ccg ttg att ccc 720
Thr Ala Ile Asp Arg Tyr Glu Leu Ile Gly Ile Gly Pro Leu Ile Pro
220 225 230
tcc gcc ttc ttg gac ggc gaa gat ccc tcc gaa acg tct tac ggc ggc 768
Ser Ala Phe Leu Asp Gly Glu Asp Pro Ser Glu Thr Ser Tyr Gly Gly
235 240 245
gat ctt ttc gaa aaa tcg gag gag aat aac tgc gtg gag tgg ttg aac 816
Asp Leu Phe Glu Lys Ser Glu Glu Asn Asn Cys Val Glu Trp Leu Asn
250 255 260
tcg aag ccg aaa tct tcg gtg gtg tat gtg tcg ttt ggg agc gtt ttg 864
Ser Lys Pro Lys Ser Ser Val Val Tyr Val Ser Phe Gly Ser Val Leu
265 270 275 280
agg ttt cca aag gca caa atg gaa gag att ggg aaa ggg cta tta gcc 912
Arg Phe Pro Lys Ala Gln Met Glu Glu Ile Gly Lys Gly Leu Leu Ala
285 290 295
tgc gga agg ccc ttt tta tgg atg ata cga gaa cag aag aat gac gac 960
Cys Gly Arg Pro Phe Leu Trp Met Ile Arg Glu Gln Lys Asn Asp Asp
300 305 310
ggc gaa gaa gaa gaa gaa gaa gaa gag ttg agt tgc att ggg gaa ttg 1008
Gly Glu Glu Glu Glu Glu Glu Glu Glu Leu Ser Cys Ile Gly Glu Leu
315 320 325
aaa aaa atg ggg aaa ata gtg tcg tgg tgc tcg cag ttg gag gtt ctg 1056
Lys Lys Met Gly Lys Ile Val Ser Trp Cys Ser Gln Leu Glu Val Leu
330 335 340
gcg cac cct gcg ttg gga tgt ttc gtg acg cat tgt ggg tgg aac tcg 1104
Ala His Pro Ala Leu Gly Cys Phe Val Thr His Cys Gly Trp Asn Ser
345 350 355 360
gct gtg gag agc ttg agt tgc ggg att ccg gtg gtg gcg gtg ccg cag 1152
Ala Val Glu Ser Leu Ser Cys Gly Ile Pro Val Val Ala Val Pro Gln
365 370 375
tgg ttt gat cag acg acg aat gcg aag ctg att gag gat gcg tgg ggg 1200
Trp Phe Asp Gln Thr Thr Asn Ala Lys Leu Ile Glu Asp Ala Trp Gly
380 385 390
aca ggg gtg aga gtg aga atg aat gaa ggg ggt ggg gtt gat gga tgt 1248
Thr Gly Val Arg Val Arg Met Asn Glu Gly Gly Gly Val Asp Gly Cys
395 400 405
gag ata gaa agg tgt gtg gag atg gtg atg gat ggg ggt gac aag acc 1296
Glu Ile Glu Arg Cys Val Glu Met Val Met Asp Gly Gly Asp Lys Thr
410 415 420
aaa cta gtg aga gaa aat gcc atc aaa tgg aag act ttg gcc aga caa 1344
Lys Leu Val Arg Glu Asn Ala Ile Lys Trp Lys Thr Leu Ala Arg Gln
425 430 435 440
gcc atg gga taggatggat cttcactcaa caatctcaac gcctttcttc 1393
Ala Met Gly
443
gtcaagttgc acacttttaa tctgctcaaa cagcggttca aataaatatc cccttccact 1453
taaaaaaaaa aaaaaaa 1470
〈210〉 3
〈211〉 2062
〈212〉 DNA
〈213〉 Verbena hybrida
〈220〉
〈221〉 CDS
〈222〉 (26)..(1408)
〈400〉 3
attttaccaa aaaaataaaa aaaaa atg agc aga gct cac gtc ctc ttg 49
Met Ser Arg Ala His Val Leu Leu
1 5
gcc aca ttc cca gca cag gga cac ata aat ccc gcc ctt caa ttc gcc 97
Ala Thr Phe Pro Ala Gln Gly His Ile Asn Pro Ala Leu Gln Phe Ala
10 15 20
aag cgt ctc gca aat gcc gac att caa gtc aca ttc ttc acc agc gtc 145
Lys Arg Leu Ala Asn Ala Asp Ile Gln Val Thr Phe Phe Thr Ser Val
25 30 35 40
tac gca tgg cgc cgc atg tcc aga acc gcc gct ggc tca aac ggg ctc 193
Tyr Ala Trp Arg Arg Met Ser Arg Thr Ala Ala Gly Ser Asn Gly Leu
45 50 55
atc aat ttt gtg tcg ttt tcc gac ggg tat gac gac ggg tta cag ccc 241
Ile Asn Phe Val Ser Phe Ser Asp Gly Tyr Asp Asp Gly Leu Gln Pro
60 65 70
gga gac gat ggg aag aac tac atg tcg gag atg aaa agc aga ggt ata 289
Gly Asp Asp Gly Lys Asn Tyr Met Ser Glu Met Lys Ser Arg Gly Ile
75 80 85
aaa gcc ttg agc gat act ctt gca gcc aat aat gtc gat caa aaa agc 337
Lys Ala Leu Ser Asp Thr Leu Ala Ala Asn Asn Val Asp Gln Lys Ser
90 95 100
agc aaa atc acg ttc gtg gtg tac tcc cac ctc ttt gca tgg gcg gcc 385
Ser Lys Ile Thr Phe Val Val Tyr Ser His Leu Phe Ala Trp Ala Ala
105 110 115 120
aag gtg gcg cgt gag ttc cat ctc cgg agc gcg cta ctc tgg att gag 433
Lys Val Ala Arg Glu Phe His Leu Arg Ser Ala Leu Leu Trp Ile Glu
125 130 135
cca gct acg gtg ttg gat ata ttt tac ttt tat ttc aac ggc tat agc 481
Pro Ala Thr Val Leu Asp Ile Phe Tyr Phe Tyr Phe Asn Gly Tyr Ser
140 145 150
gac gaa atc gat gcg ggt tcg gat gct att cac ttg ccc gga gga ctc 529
Asp Glu Ile Asp Ala Gly Ser Asp Ala Ile His Leu Pro Gly Gly Leu
155 160 165
cca gtg ctg gcc cag cgt gat tta ccg tct ttc ctt ctt cct tcc acg 577
Pro Val Leu Ala Gln Arg Asp Leu Pro Ser Phe Leu Leu Pro Ser Thr
170 175 180
cat gag aga ttc cgt tca ctg atg aag gag aaa ttg gaa act tta gaa 625
His Glu Arg Phe Arg Ser Leu Met Lys Glu Lys Leu Glu Thr Leu Glu
185 190 195 200
ggt gaa gaa aaa cct aag gtc ttg gtg aac agc ttt gat gcg ttg gag 673
Gly Glu Glu Lys Pro Lys Val Leu Val Asn Ser Phe Asp Ala Leu Glu
205 210 215
cct gat gcg ctc aag gcc att gat aag tac gag atg att gca atc ggg 721
Pro Asp Ala Leu Lys Ala Ile Asp Lys Tyr Glu Met Ile Ala Ile Gly
220 225 230
ccg ttg att cct tcc gca ttc ttg gac ggt aaa gat cct tcg gac agg 769
Pro Leu Ile Pro Ser Ala Phe Leu Asp Gly Lys Asp Pro Ser Asp Arg
235 240 245
tct ttc ggc gga gat ttg ttc gag aaa ggg tcg aat gac gac gat tgc 817
Ser Phe Gly Gly Asp Leu Phe Glu Lys Gly Ser Asn Asp Asp Asp Cys
250 255 260
ctc gaa tgg ttg agc acg aat cct cga tct tcg gtg gtt tac gtt tcg 865
Leu Glu Trp Leu Ser Thr Asn Pro Arg Ser Ser Val Val Tyr Val Ser
265 270 275 280
ttc gga agc ttc gtt aat acg acg aag tcg caa atg gaa gag ata gca 913
Phe Gly Ser Phe Val Asn Thr Thr Lys Ser Gln Met Glu Glu Ile Ala
285 290 295
aga ggg ctg tta gat tgt ggg agg ccg ttt ttg tgg gtg gta aga gta 961
Arg Gly Leu Leu Asp Cys Gly Arg Pro Phe Leu Trp Val Val Arg Val
300 305 310
aac gaa gga gaa gag gta ttg ata agt tgc atg gag gag ttg aaa cga 1009
Asn Glu Gly Glu Glu Val Leu Ile Ser Cys Met Glu Glu Leu Lys Arg
315 320 325
gtg ggg aaa att gta tct tgg tgt tct caa ttg gaa gtc ctg acg cat 1057
Val Gly Lys Ile Val Ser Trp Cys Ser Gln Leu Glu Val Leu Thr His
330 335 340
ccc tcg ttg gga tgt ttc gtg aca cac tgc ggg tgg aat tcg act cta 1105
Pro Ser Leu Gly Cys Phe Val Thr His Cys Gly Trp Asn Ser Thr Leu
345 350 355 360
gag agt ata tct ttc ggg gtt ccg atg gtg gct ttt ccg cag tgg ttc 1153
Glu Ser Ile Ser Phe Gly Val Pro Met Val Ala Phe Pro Gln Trp Phe
365 370 375
gat caa ggg acg aat gcg aag ctg atg gag gat gtg tgg agg acg ggt 1201
Asp Gln Gly Thr Asn Ala Lys Leu Met Glu Asp Val Trp Arg Thr Gly
380 385 390
gtg aga gtc aga gct aat gag gag gct agc gtc gtt gat ggt gat gaa 1249
Val Arg Val Arg Ala Asn Glu Glu Ala Ser Val Val Asp Gly Asp Glu
395 400 405
att agg aga tgt att gag gag gtt atg gat ggg gga gaa aag agt agg 1297
Ile Arg Arg Cys Ile Glu Glu Val Met Asp Gly Gly Glu Lys Ser Arg
410 415 420
aaa ctt aga gag agt gct ggc aag tgg aag gat ttg gca aga aaa gct 1345
Lys Leu Arg Glu Ser Ala Gly Lys Trp Lys Asp Leu Ala Arg Lys Ala
425 430 435 440
atg gag gaa gat gga tct tca gtt aac aac ctc aag gtc ttt ctt gat 1393
Met Glu Glu Asp Gly Ser Ser Val Asn Asn Leu Lys Val Phe Leu Asp
445 450 455
gag gtt gta ggt atc ta aagacgtaaa tgaggtcccc ataggcaaaa 1440
Glu Val Val Gly Ile
460
ttgcaaattt catctcgtaa gttgaatact ttttggcttt aattttgttc gagtttgttt 1500
ttcaaaattt atcttgtaat tttacattga gtgtaaattt agtctgattt taactggaaa 1560
aatataaaat tcattgttga gactcttcat caaaatcatc tgatttcctt tattgtcttg 1620
gtcaaaattc tcatatcaat tggaaaaaat aaatttcaaa atcgtccaat tttgaaccaa 1680
gaaagaagta taatttgacc aaaataataa aaggattcaa gtgatcttga tgaagtgtct 1740
gagcgacgag ttctatattt ttccaccgaa tttctaacga gtttttgaat tttttttagc 1800
caaaatcgga ctaactttgt acaaaatgaa aagttatatg atgaaatttt aaaaaacaaa 1860
ctcagacaat aataaagccc gaaagtagta aaattacctg acgaaatttg caatttcgcc 1920
tcctatttta atttttttgg tgtgtttaat aaatcggtta ttttactttt aattaaaata 1980
aaagtgagat gcatgatagc ttggtgagta tatatgagtt gatggtaatg tacgatattt 2040
tctaaaaaaa aaaaaaaaaa aa 2062
〈210〉 4
〈211〉 1671
〈212〉 DNA
〈213〉 Torenia hybrira
〈220〉
〈221〉 CDS
〈222〉 (45)..(1478)
〈223〉 Xaa (64) is Cys or Phe, Xaa(65) is Ser or Pro
〈400〉 4
aacacataaa aaaaaaataa aagaagaaat aattaaaaaa aaaa atg gtt aac 53
Met Val Asn
1
aaa cgc cat att cta cta gca aca ttc cca gca caa ggc cac ata aac 101
Lys Arg His Ile Leu Leu Ala Thr Phe Pro Ala Gln Gly His Ile Asn
5 10 15
cct tct ctc gag ttc gcc aaa agg ctc ctc aac acc gga tac gtc gac 149
Pro Ser Leu Glu Phe Ala Lys Arg Leu Leu Asn Thr Gly Tyr Val Asp
20 25 30 35
caa gtc aca ttc ttc acg agt gta tac gca ttg aga cgc atg cgc ttc 197
Gln Val Thr Phe Phe Thr Ser Val Tyr Ala Leu Arg Arg Met Arg Phe
40 45 50
gaa acc gat ccg agc agc aga atc gat ttc gtg gca tkt yca gat tct 245
Glu Thr Asp Pro Ser Ser Arg Ile Asp Phe Val Ala Xaa Xaa Asp Ser
55 60 65
tac gat gat ggc tta aag aaa ggc gac gat ggc aaa aac tac atg tcg 293
Tyr Asp Asp Gly Leu Lys Lys Gly Asp Asp Gly Lys Asn Tyr Met Ser
70 75 80
gag atg aga aag cgc gga acg aag gcc tta aag gac act ctt att aag 341
Glu Met Arg Lys Arg Gly Thr Lys Ala Leu Lys Asp Thr Leu Ile Lys
85 90 95
ctc aac gat gct gcg atg gga agt gaa tgt tac aat cgc gtg agc ttt 389
Leu Asn Asp Ala Ala Met Gly Ser Glu Cys Tyr Asn Arg Val Ser Phe
100 105 110 115
gtg gtg tac tct cat cta ttt tcg tgg gca gct gaa gtg gcc cgt gaa 437
Val Val Tyr Ser His Leu Phe Ser Trp Ala Ala Glu Val Ala Arg Glu
120 125 130
gtc gac gtg ccg agt gcc ctt ctt tgg att gaa ccg gct acg gtt ttc 485
Val Asp Val Pro Ser Ala Leu Leu Trp Ile Glu Pro Ala Thr Val Phe
135 140 145
gat gtg tac tat ttt tac ttc aat ggg tat gcc gat gat atc gat gcg 533
Asp Val Tyr Tyr Phe Tyr Phe Asn Gly Tyr Ala Asp Asp Ile Asp Ala
150 155 160
ggc tca gat caa atc caa ctg ccc aat ctt ccg cag ctc tcc aag caa 581
Gly Ser Asp Gln Ile Gln Leu Pro Asn Leu Pro Gln Leu Ser Lys Gln
165 170 175
gat ctc ccc tct ttc cta ctc cct tcg agc ccc gcg aga ttc cga acc 629
Asp Leu Pro Ser Phe Leu Leu Pro Ser Ser Pro Ala Arg Phe Arg Thr
180 185 190 195
cta atg aaa gaa aag ttc gac acg ctc gac aaa gaa ccg aaa gcg aag 677
Leu Met Lys Glu Lys Phe Asp Thr Leu Asp Lys Glu Pro Lys Ala Lys
200 205 210
gtc ttg ata aac acg ttc gac gca tta gaa acc gaa caa ctc aaa gcc 725
Val Leu Ile Asn Thr Phe Asp Ala Leu Glu Thr Glu Gln Leu Lys Ala
215 220 225
atc gac agg tat gaa cta ata tcc atc ggc cca tta atc cca tca tcg 773
Ile Asp Arg Tyr Glu Leu Ile Ser Ile Gly Pro Leu Ile Pro Ser Ser
230 235 240
ata ttc tca gat ggc aac gac ccc tca tca agc aac aaa tcc tac ggt 821
Ile Phe Ser Asp Gly Asn Asp Pro Ser Ser Ser Asn Lys Ser Tyr Gly
245 250 255
gga gac ctc ttc aga aaa gcc gat gaa act tac atg gac tgg cta aac 869
Gly Asp Leu Phe Arg Lys Ala Asp Glu Thr Tyr Met Asp Trp Leu Asn
260 265 270 275
tca aaa ccc gaa tca tcg gtc gtt tac gtt tcg ttc ggg agc ctc ctg 917
Ser Lys Pro Glu Ser Ser Val Val Tyr Val Ser Phe Gly Ser Leu Leu
280 285 290
agg ctc ccg aaa ccc caa atg gaa gaa ata gca ata ggg ctt tca gac 965
Arg Leu Pro Lys Pro Gln Met Glu Glu Ile Ala Ile Gly Leu Ser Asp
295 300 305
acc aaa tcg cca gtt ctc tgg gtg ata aga aga aac gaa gag ggc gac 1013
Thr Lys Ser Pro Val Leu Trp Val Ile Arg Arg Asn Glu Glu Gly Asp
310 315 320
gaa caa gag caa gca gaa gaa gaa gag aag ctg ctg agc ttc ttt gat 1061
Glu Gln Glu Gln Ala Glu Glu Glu Glu Lys Leu Leu Ser Phe Phe Asp
325 330 335
cgt cac gga act gaa cga ctc ggg aaa atc gtg aca tgg tgc tca caa 1109
Arg His Gly Thr Glu Arg Leu Gly Lys Ile Val Thr Trp Cys Ser Gln
340 345 350 355
ttg gat gtt ctg acg cat aag tcg gtg gga tgc ttc gtg acg cat tgc 1157
Leu Asp Val Leu Thr His Lys Ser Val Gly Cys Phe Val Thr His Cys
360 365 370
ggt tgg aat tct gct atc gag agc ctg gct tgt ggt gtg ccc gtg gtg 1205
Gly Trp Asn Ser Ala Ile Glu Ser Leu Ala Cys Gly Val Pro Val Val
375 380 385
tgc ttt cct caa tgg ttc gat caa ggg act aat gcg aag atg atc gaa 1253
Cys Phe Pro Gln Trp Phe Asp Gln Gly Thr Asn Ala Lys Met Ile Glu
390 395 400
gat gtg tgg agg agt ggt gtg aga gtc aga gtg aat gag gaa ggc ggc 1301
Asp Val Trp Arg Ser Gly Val Arg Val Arg Val Asn Glu Glu Gly Gly
405 410 415
gtt gtt gat agg cgt gag att aag agg tgc gtc tcg gag gtt ata aag 1349
Val Val Asp Arg Arg Glu Ile Lys Arg Cys Val Ser Glu Val Ile Lys
420 425 430 435
agt cga gag ttg aga gaa agc gca atg atg tgg aag ggt ttg gct aaa 1397
Ser Arg Glu Leu Arg Glu Ser Ala Met Met Trp Lys Gly Leu Ala Lys
440 445 450
gaa gct atg gat gaa gaa cgt gga tca tca atg aac aat ctg aag aat 1445
Glu Ala Met Asp Glu Glu Arg Gly Ser Ser Met Asn Asn Leu Lys Asn
455 460 465
ttt att act agg att att aat gaa aat gcc tca ta agttgtacta 1490
Phe Ile Thr Arg Ile Ile Asn Glu Asn Ala Ser
470 475
tatatgttat tattgttgtt atggacgtcg aattaagtat tagttaaatg atatgtattt 1550
agaggaaggc caaaacgggc tacacccggc aggccacggg ttggaaaagc ccgccatgat 1610
ttaaaatata tattttaaaa taaatatttt ctactattaa actaaaaaaa aaaaaaaaaa 1670
a 1671
〈210〉 5
〈211〉 1437
〈212〉 DNA
〈213〉 Perilla frutescens
〈220〉
〈221〉 CDS
〈222〉 (294)..(1298)
〈400〉 5
ttcaaaactc ataacgtgat tgagctaatg tgcacatctt cctcttcaaa gtctacagtg 60
tcatcctacc agcatcatca tgatcaatct ctttataatg aggagaatgg agtaacaagg 120
agtgggtttt gttactcagc ttcaacctac gtacgtacta ctactgactc aactctcaag 180
agaatgaata taatatataa tgggcgatag atctttgtag atatgtaggt gtagcctgca 240
ggtggttaat taatttccgg tgtgggaaaa taaataaata aataaatata gcg 293
atg agc agc agc agc agc aga agg tgg aga gag aat gag ggg atg cga 341
Met Ser Ser Ser Ser Ser Arg Arg Trp Arg Glu Asn Glu Gly Met Arg
1 5 10 15
agg aca ttg ctg ggg ttg ggt ttg ggg cag ttg gtt tct ttc gat ttg 389
Arg Thr Leu Leu Gly Leu Gly Leu Gly Gln Leu Val Ser Phe Asp Leu
20 25 30
gct atc atg acc ttt tct gct tct ttg gtt tca acc aca gtg gat gca 437
Ala Ile Met Thr Phe Ser Ala Ser Leu Val Ser Thr Thr Val Asp Ala
35 40 45
cca ctt act atg tcc ttc act aca tac act gtt gtg gcc ctg ctc tat 485
Pro Leu Thr Met Ser Phe Thr Thr Tyr Thr Val Val Ala Leu Leu Tyr
50 55 60
gga acc atc ttg ctt tac cgc cgc cac aaa ttc ttg gtt cca tgg tac 533
Gly Thr Ile Leu Leu Tyr Arg Arg His Lys Phe Leu Val Pro Trp Tyr
65 70 75 80
tgg tat gct ctc ctg ggc ttc gtg cac ctc cac ggc aat tat ctt gtt 581
Trp Tyr Ala Leu Leu Gly Phe Val His Leu His Gly Asn Tyr Leu Val
85 90 95
aat aaa gca ttc gag ttg aca tcg att acc agt ctc agc ata ctg gat 629
Asn Lys Ala Phe Glu Leu Thr Ser Ile Thr Ser Leu Ser Ile Leu Asp
100 105 110
tgt tgc aca atc ctc tcg tcc atc atc ttt aca tgg atc ttc cta ccc 677
Cys Cys Thr Ile Leu Ser Ser Ile Ile Phe Thr Trp Ile Phe Leu Pro
115 120 125
aca aaa tac tct cta tac cac ttt ctc ggt gct gct att tct cta cga 725
Thr Lys Tyr Ser Leu Tyr His Phe Leu Gly Ala Ala Ile Ser Leu Arg
130 135 140
ggc ctc ctc ctc gtc ctt ctt tcc gac tca ggg gtc act gct gct ggt 773
Gly Leu Leu Leu Val Leu Leu Ser Asp Ser Gly Val Thr Ala Ala Gly
145 150 155 160
tcg aat cct ctt ttg ggt gat ttt ctt gtc ata aca ggc tct att ttg 821
Ser Asn Pro Leu Leu Gly Asp Phe Val Val Ile Thr Gly Ser Ile Leu
165 170 175
ttc aca ctc agc act gtt ggt cag gaa tac tgc gtg aag agg aaa gat 869
Phe Thr Leu Ser Thr Val Gly Gln Glu Tyr Cys Val Lys Arg Lys Asp
180 185 190
cgt att gaa gta gta gca atg atc ggt gta ttt ggt atg ctc atc agt 917
Arg Ile Glu Val Val Ala Met Ile Gly Val Phe Gly Met Leu Ile Ser
195 200 205
gca acc gag att act gtg ctg gag agg aat gcc ctc tca tca atg cag 965
Ala Thr Glu Ile Thr Val Leu Glu Arg Asn Ala Leu Ser Ser Met Gln
210 215 220
tgg tct act gga ctt ttg gca gcc tat gtt gtt tat gca ctg tcc agc 1013
Trp Ser Thr Gly Leu Leu Ala Ala Tyr Val Val Tyr Ala Leu Ser Ser
225 230 235 240
ttc ctc ttc tgc aca ctc acc cct ttt ctt ctc aag atg agt ggc gct 1061
Phe Leu Phe Cys Thr Leu Thr Pro Phe Leu Leu Lys Met Ser Gly Ala
245 250 255
gca ttt ttc aat ctt tcc atg ctt aca tct gat atg tgg gct gtt gca 1109
Ala Phe Phe Asn Leu Ser Met Leu Thr Ser Asp Met Trp Ala Val Ala
260 265 270
att agg aca ttc ata tac aac cag gag gtt gat tgg tta tac tat ttg 1157
Ile Arg Thr Phe Ile Tyr Asn Gln Glu Val Asp Trp Leu Tyr Tyr Leu
275 280 285
gcc ttt tgt ctc gtt gtt gtt gga ata ttc ata tat aca aaa aca gag 1205
Ala Phe Cys Leu Val Val Val Gly Ile Phe Ile Tyr Thr Lys Thr Glu
290 295 300
aag gat cct aac aat acg aga gcc ctt gag aat gga aac ttg gat cat 1253
Lys Asp Pro Asn Asn Thr Arg Ala Leu Glu Asn Gly Asn Leu Asp His
305 310 315 320
gaa tat agt ctc ctt gag gat caa gat gac aca cca aga aaa cca ta 1300
Glu Tyr Ser Leu Leu Glu Asp Gln Asp Asp Thr Pro Arg Lys Pro
325 330 335
gctagctttg cccacaatct tttcatcaac agttttaaat aattcgtgag ggggagagag 1360
atcgagatac taattaatgg acgtctatta tatagttgga ggtttttgtt ttatttattt 1420
atttgagtaa aaaaaaa 1437
〈210〉 6
〈211〉 2105
〈212〉 DNA
〈213〉 Petunia hybrida
〈220〉
〈221〉 CDS
〈222〉 (341)..(1744)
〈400〉 6
agtgagcgca acgcaattaa tgtgagttag ctcactcatt aggcacccca ggctttacac 60
tttatgcttc cggctcgtat gttgtgtgga attgtgagcg gataacaatt tcacacagga 120
aacagctatg accatgatta cgccaagctc gaaattaacc ctcactaaag ggaacaaaag 180
ctggagctcc acgcggtggc ggccgctcta gaactagtgg atcccccggg ctgcaggaat 240
tccgttgctg tcgccacaat ttacaaacca agaaattaag catccctttc ccccccttaa 300
aaaacataca agtttttaat ttttcactaa gcaagaaaat atg gtg cag cct cat 355
Met Val Gln Pro His
1 5
gtc atc tta aca aca ttt cca gca caa ggc cat att aat cca gca ctt 403
Val Ile Leu Thr Thr Phe Pro Ala Gln Gly His Ile Asn Pro Ala Leu
10 15 20
caa ttt gcc aag aat ctt gtc aag atg ggc ata gaa gtg aca ttt tct 451
Gln Phe Ala Lys Asn Leu Val Lys Met Gly Ile Glu Val Thr Phe Ser
25 30 35
aca agc att tat gcc caa agc cgt atg gat gaa aaa tcc att ctt aat 499
Thr Ser Ile Tyr Ala Gln Ser Arg Met Asp Glu Lys Ser Ile Leu Asn
40 45 50
gca cca aaa gga ttg aat ttc att cca ttt tcc gat ggc ttt gat gaa 547
Ala Pro Lys Gly Leu Asn Phe Ile Pro Phe Ser Asp Gly Phe Asp Glu
55 60 65
ggt ttt gat cat tca aaa gac cct gta ttt tac atg tca caa ctt cgt 595
Gly Phe Asp His Ser Lys Asp Pro Val Phe Tyr Met Ser Gln Leu Arg
70 75 80 85
aaa tgt gga agt gaa act gtc aaa aaa ata att ctc act tgc tct gaa 643
Lys Cys Gly Ser Glu Thr Val Lys Lys Ile Ile Leu Thr Cys Ser Glu
90 95 100
aat gga cag cct ata act tgc cta ctt tac tcc att ttc ctt cct tgg 691
Asn Gly Gln Pro Ile Thr Cys Leu Leu Tyr Ser Ile Phe Leu Pro Trp
105 110 115
gca gca gag gta gca cgt gaa gtt cac atc cct tct gct ctt ctt tgg 739
Ala Ala Glu Val Ala Arg Glu Val His Ile Pro Ser Ala Leu Leu Trp
120 125 130
agt caa cca gca aca ata ttg gac ata tat tac ttc aac ttt cat gga 787
Ser Gln Pro Ala Thr Ile Leu Asp Ile Tyr Tyr Phe Asn Phe His Gly
135 140 145
tat gaa aaa gct atc gct aat gaa tcc aat gat cca aat tgg tcc att 835
Tyr Glu Lys Ala Ile Ala Asn Glu Ser Asn Asp Pro Asn Trp Ser Ile
150 155 160 165
caa ctt ccc ggg ctt cca cta ctg gaa act cga gat ctt cct tca ttt 883
Gln Leu Pro Gly Leu Pro Leu Leu Glu Thr Arg Asp Leu Pro Ser Phe
170 175 180
tta ctt cct tat ggt gca aaa ggc agt ctt cga gtt gca ctt cca cca 931
Leu Leu Pro Tyr Gly Ala Lys Gly Ser Leu Arg Val Ala Leu Pro Pro
185 190 195
ttc aaa gaa ttg ata gac aca tta gat gct gaa acc act cct aag att 979
Phe Lys Glu Leu Ile Asp Thr Leu Asp Ala Glu Thr Thr Pro Lys Ile
200 205 210
ctt gtg aat aca ttt gat gaa tta gag cct gag gca ctc aat gca att 1027
Leu Val Asn Thr Phe Asp Glu Leu Glu Pro Glu Ala Leu Asn Ala Ile
215 220 225
gaa ggt tat aag ttt tat gga att gga ccg ttg att cct tct gct ttc 1075
Glu Gly Tyr Lys Phe Tyr Gly Ile Gly Pro Leu Ile Pro Ser Ala Phe
230 235 240 245
ttg ggt gga aat gac cct tta gat gct tca ttt ggt ggt gat ctt ttt 1123
Leu Gly Gly Asn Asp Pro Leu Asp Ala Ser Phe Gly Gly Asp Leu Phe
250 255 260
caa aat tca aat gac tat atg gaa tgg tta aac tca aag cca aat tca 1171
Gln Asn Ser Asn Asp Tyr Met Glu Trp Leu Asn Ser Lys Pro Asn Ser
265 270 275
tca ctt gtt tat ata tct ttt ggg agt cta atg aat cca tct att agc 1219
Ser Leu Val Tyr Ile Ser Phe Gly Ser Leu Met Asn Pro Ser Ile Ser
280 285 290
caa atg gag gag ata tca aaa ggg ttg ata gac ata gga agg ccg ttt 1267
Gln Met Glu Glu Ile Ser Lys Gly Leu Ile Asp Ile Gly Arg Pro Phe
295 300 305
tta tgg gtg ata aaa gaa aat gaa aaa ggc aaa gaa gaa gag aat aaa 1315
Leu Trp Val Ile Lys Glu Asn Glu Lys Gly Lys Glu Glu Glu Asn Lys
310 315 320 325
aag ctt ggt tgt att gaa gaa ttg gaa aaa ata gga aaa ata gtt cca 1363
Lys Leu Gly Cys Ile Glu Glu Leu Glu Lys Ile Gly Lys Ile Val Pro
330 335 340
tgg tgt tca caa ctt gaa gtt cta aaa cat cca tct tta gga tgt ttt 1411
Trp Cys Ser Gln Leu Glu Val Leu Lys His Pro Ser Leu Gly Cys Phe
345 350 355
gtt tct cat tgt gga tgg aat tca gcc tta gag agt tta gct tgt gga 1459
Val Ser His Cys Gly Trp Asn Ser Ala Leu Glu Ser Leu Ala Cys Gly
360 365 370
gtg cca gtt gtg gca ttt cct caa tgg aca gat caa atg aca aat gcc 1507
Val Pro Val Val Ala Phe Pro Gln Trp Thr Asp Gln Met Thr Asn Ala
375 380 385
aaa caa gtt gaa gat gtc tgg aaa agt gga gta aga gtg aga ata aat 1555
Lys Gln Val Glu Asp Val Trp Lys Ser Gly Val Arg Val Arg Ile Asn
390 395 400 405
gaa gat ggt gtt gtt gaa agt gag gaa atc aaa agg tgt att gaa ttg 1603
Glu Asp Gly Val Val Glu Ser Glu Glu Ile Lys Arg Cys Ile Glu Leu
410 415 420
gta atg gat gga gga gag aaa ggg gaa gaa ttg aga aag aat gct aag 1651
Val Met Asp Gly Gly Glu Lys Gly Glu Glu Leu Arg Lys Asn Ala Lys
425 430 435
aaa tgg aaa gaa ttg gct aga gaa gct gtg aag gaa ggt gga tct tca 1699
Lys Trp Lys Glu Leu Ala Arg Glu Ala Val Lys Glu Gly Gly Ser Ser
440 445 450
cac aag aat tta aag gct ttt att gat gat gtt gcc aaa ggg ttt 1744
His Lys Asn Leu Lys Ala Phe Ile Asp Asp Val Ala Lys Gly Phe
455 460 465
taatat ttacaggctt ttgccgtgat attacttccc ctagttggcg attcactctt 1800
tgtggacttg cttgacaaaa aactgaggga atgtgctaag acacgctaat gctttaagaa 1860
gtcatttcca aggcttgaag cctgctttta aaacttatta gccagtaatc tatagggttc 1920
tcttctattt ttctctgtct ctctttttag cctttttctt tccaaggttt aagaatagcg 1980
tgaacatagc ttagtacgta gtcttggtat ctctatctta ccaagtgcaa gattatgctt 2040
atgctgtcct cctaaatttc ttaataaaat gcaagatgaa aaagtacaaa aaaaaaaaaa 2100
aaaaa 2105
〈210〉 7
〈211〉 460
〈212〉 PRT
〈213〉 Perilla frutescens
〈400〉 7
Met Val Arg Arg Arg Val Leu Leu Ala Thr Phe Pro Ala Gln Gly His
1 5 10 15
Ile Asn Pro Ala Leu Gln Phe Ala Lys Arg Leu Leu Lys Ala Gly Thr
20 25 30
Asp Val Thr Phe Phe Thr Ser Val Tyr Ala Trp Arg Arg Met Ala Asn
35 40 45
Thr Ala Ser Ala Ala Ala Gly Asn Pro Pro Gly Leu Asp Phe Val Ala
50 55 60
Phe Ser Asp Gly Tyr Asp Asp Gly Leu Lys Pro Cys Gly Asp Gly Lys
65 70 75 80
Arg Tyr Met Ser Glu Met Lys Ala Arg Gly Ser Glu Ala Leu Arg Asn
85 90 95
Leu Leu Leu Asn Asn His Asp Val Thr Phe Val Val Tyr Ser His Leu
100 105 110
Phe Ala Trp Ala Ala Glu Val Ala Arg Glu Ser Gln Val Pro Ser Ala
115 120 125
Leu Leu Trp Val Glu Pro Ala Thr Val Leu Cys Ile Tyr Tyr Phe Tyr
130 135 140
Phe Asn Gly Tyr Ala Asp Glu Ile Asp Ala Gly Ser Asp Glu Ile Gln
145 150 155 160
Leu Pro Arg Leu Pro Pro Leu Glu Gln Arg Ser Leu Pro Thr Phe Leu
165 170 175
Leu Pro Glu Thr Pro Glu Arg Phe Arg Leu Met Met Lys Glu Lys Leu
180 185 190
Glu Thr Leu Asp Gly Glu Glu Lys Ala Lys Val Leu Val Asn Thr Phe
195 200 205
Asp Ala Leu Glu Pro Asp Ala Leu Thr Ala Ile Asp Arg Tyr Glu Leu
210 215 220
Ile Gly Ile Gly Pro Leu Ile Pro Ser Ala Phe Leu Asp Gly Gly Asp
225 230 235 240
Pro Ser Glu Thr Ser Tyr Gly Gly Asp Leu Phe Glu Lys Ser Glu Glu
245 250 255
Asn Asn Cys Val Glu Trp Leu Asp Thr Lys Pro Lys Ser Ser Val Val
260 265 270
Tyr Val Ser Phe Gly Ser Val Leu Arg Phe Pro Lys Ala Gln Met Glu
275 280 285
Glu Ile Gly Lys Gly Leu Leu Ala Cys Gly Arg Pro Phe Leu Trp Met
290 295 300
Ile Arg Glu Gln Lys Asn Asp Asp Gly Glu Glu Glu Glu Glu Glu Leu
305 310 315 320
Ser Cys Ile Gly Glu Leu Lys Lys Met Gly Lys Ile Val Ser Trp Cys
325 330 335
Ser Gln Leu Glu Val Leu Ala His Pro Ala Leu Gly Cys Phe Val Thr
340 345 350
His Cys Gly Trp Asn Ser Ala Val Glu Ser Leu Ser Cys Gly Val Pro
355 360 365
Val Val Ala Val Pro Gln Trp Phe Asp Gln Thr Thr Asn Ala Lys Leu
370 375 380
Ile Glu Asp Ala Trp Gly Thr Gly Val Arg Val Arg Met Asn Glu Gly
385 390 395 400
Gly Gly Val Asp Gly Ser Glu Ile Glu Arg Cys Val Glu Met Val Met
405 410 415
Asp Gly Gly Glu Lys Ser Lys Leu Val Arg Glu Asn Ala Ile Lys Trp
420 425 430
Lys Thr Leu Ala Arg Glu Ala Met Gly Glu Asp Gly Ser Ser Leu Lys
435 440 445
Asn Leu Asn Ala Phe Leu His Gln Val Ala Arg Ala
450 455 460
〈210〉 8
〈211〉 443
〈212〉 PRT
〈213〉 Perilla frutescens
〈400〉 8
Met Val Arg Arg Arg Val Leu Leu Ala Thr Phe Pro Ala Gln Gly His
1 5 10 15
Ile Asn Pro Ala Leu Gln Phe Ala Lys Arg Leu Leu Lys Ala Gly Thr
20 25 30
Asp Val Thr Phe Phe Thr Ser Val Tyr Ala Trp Arg Arg Met Ala Asn
35 40 45
Thr Ala Ser Ala Ala Ala Gly Asn Pro Pro Gly Leu Asp Phe Val Ala
50 55 60
Phe Ser Asp Gly Tyr Asp Asp Gly Leu Lys Pro Gly Gly Asp Gly Lys
65 70 75 80
Arg Tyr Met Ser Glu Met Lys Ala Arg Gly Ser Glu Ala Leu Arg Asn
85 90 95
Leu Leu Leu Asn Asn Asp Asp Val Thr Phe Val Val Tyr Ser His Leu
100 105 110
Phe Ala Trp Ala Ala Glu Val Ala Arg Leu Ser His Val Pro Thr Ala
115 120 125
Leu Leu Trp Val Glu Pro Ala Thr Val Leu Cys Ile Tyr His Phe Tyr
130 135 140
Phe Asn Gly Tyr Ala Asp Glu Ile Asp Ala Gly Ser Asn Glu Ile Gln
145 150 155 160
Leu Pro Arg Leu Pro Ser Leu Glu Gln Arg Ser Leu Pro Thr Phe Leu
165 170 175
Leu Pro Ala Thr Pro Glu Arg Phe Arg Leu Met Met Lys Glu Lys Leu
180 185 190
Glu Thr Leu Asp Gly Glu Glu Lys Ala Lys Val Leu Val Asn Thr Phe
195 200 205
Asp Ala Leu Glu Pro Asp Ala Leu Thr Ala Ile Asp Arg Tyr Glu Leu
210 215 220
Ile Gly Ile Gly Pro Leu Ile Pro Ser Ala Phe Leu Asp Gly Glu Asp
225 230 235 240
Pro Ser Glu Thr Ser Tyr Gly Gly Asp Leu Phe Glu Lys Ser Glu Glu
245 250 255
Asn Asn Cys Val Glu Trp Leu Asn Ser Lys Pro Lys Ser Ser Val Val
260 265 270
Tyr Val Ser Phe Gly Ser Val Leu Arg Phe Pro Lys Ala Gln Met Glu
275 280 285
Glu Ile Gly Lys Gly Leu Leu Ala Cys Gly Arg Pro Phe Leu Trp Met
290 295 300
Ile Arg Glu Gln Lys Asn Asp Asp Gly Glu Glu Glu Glu Glu Glu Glu
305 310 315 320
Glu Leu Ser Cys Ile Gly Glu Leu Lys Lys Met Gly Lys Ile Val Ser
325 330 335
Trp Cys Ser Gln Leu Glu Val Leu Ala His Pro Ala Leu Gly Cys Phe
340 345 350
Val Thr His Cys Gly Trp Asn Ser Ala Val Glu Ser Leu Ser Cys Gly
355 360 365
Ile Pro Val Val Ala Val Pro Gln Trp Phe Asp Gln Thr Thr Asn Ala
370 375 380
Lys Leu Ile Glu Asp Ala Trp Gly Thr Gly Val Arg Val Arg Met Asn
385 390 395 400
Glu Gly Gly Gly Val Asp Gly Cys Glu Ile Glu Arg Cys Val Glu Met
405 410 415
Val Met Asp Gly Gly Asp Lys Thr Lys Leu Val Arg Glu Asn Ala Ile
420 425 430
Lys Trp Lys Thr Leu Ala Arg Gln Ala Met Gly
435 440
〈210〉 9
〈211〉 461
〈212〉 PRT
〈213〉 Verbena hybrida
〈400〉 9
Met Ser Arg Ala His Val Leu Leu Ala Thr Phe Pro Ala Gln Gly His
1 5 10 15
Ile Asn Pro Ala Leu Gln Phe Ala Lys Arg Leu Ala Asn Ala Asp Ile
20 25 30
Gln Val Thr Phe Phe Thr Ser Val Tyr Ala Trp Arg Arg Met Ser Arg
35 40 45
Thr Ala Ala Gly Ser Asn Gly Leu Ile Asn Phe Val Ser Phe Ser Asp
50 55 60
Gly Tyr Asp Asp Gly Leu Gln Pro Gly Asp Asp Gly Lys Asn Tyr Met
65 70 75 80
Ser Glu Met Lys Ser Arg Gly Ile Lys Ala Leu Ser Asp Thr Leu Ala
85 90 95
Ala Asn Asn Val Asp Gln Lys Ser Ser Lys Ile Thr Phe Val Val Tyr
100 105 110
Ser His Leu Phe Ala Trp Ala Ala Lys Val Ala Arg Glu Phe His Leu
115 120 125
Arg Ser Ala Leu Leu Trp Ile Glu Pro Ala Thr Val Leu Asp Ile Phe
130 135 140
Tyr Phe Tyr Phe Asn Gly Tyr Ser Asp Glu Ile Asp Ala Gly Ser Asp
145 150 155 160
Ala Ile His Leu Pro Gly Gly Leu Pro Val Leu Ala Gln Arg Asp Leu
165 170 175
Pro Ser Phe Leu Leu Pro Ser Thr His Glu Arg Phe Arg Ser Leu Met
180 185 190
Lys Glu Lys Leu Glu Thr Leu Glu Gly Glu Glu Lys Pro Lys Val Leu
195 200 205
Val Asn Ser Phe Asp Ala Leu Glu Pro Asp Ala Leu Lys Ala Ile Asp
210 215 220
Lys Tyr Glu Met Ile Ala Ile Gly Pro Leu Ile Pro Ser Ala Phe Leu
225 230 235 240
Asp Gly Lys Asp Pro Ser Asp Arg Ser Phe Gly Gly Asp Leu Phe Glu
245 250 255
Lys Gly Ser Asn Asp Asp Asp Cys Leu Glu Trp Leu Ser Thr Asn Pro
260 265 270
Arg Ser Ser Val Val Tyr Val Ser Phe Gly Ser Phe Val Asn Thr Thr
275 280 285
Lys Ser Gln Met Glu Glu Ile Ala Arg Gly Leu Leu Asp Cys Gly Arg
290 295 300
Pro Phe Leu Trp Val Val Arg Val Asn Glu Gly Glu Glu Val Leu Ile
305 310 315 320
Ser Cys Met Glu Glu Leu Lys Arg Val Gly Lys Ile Val Ser Trp Cys
325 330 335
Ser Gln Leu Glu Val Leu Thr His Pro Ser Leu Gly Cys Phe Val Thr
340 345 350
His Cys Gly Trp Asn Ser Thr Leu Glu Ser Ile Ser Phe Gly Val Pro
355 360 365
Met Val Ala Phe Pro Gln Trp Phe Asp Gln Gly Thr Asn Ala Lys Leu
370 375 380
Met Glu Asp Val Trp Arg Thr Gly Val Arg Val Arg Ala Asn Glu Glu
385 390 395 400
Gly Ser Val Val Asp Gly Asp Glu Ile Arg Arg Cys Ile Glu Glu Val
405 410 415
Met Asp Gly Gly Glu Lys Ser Arg Lys Leu Arg Glu Ser Ala Gly Lys
420 425 430
Trp Lys Asp Leu Ala Arg Lys Ala Met Glu Glu Asp Gly Ser Ser Val
435 440 445
Asn Asn Leu Lys Val Phe Leu Asp Glu Val Val Gly Ile
450 455 460
〈210〉 10
〈211〉 478
〈212〉 PRT
〈213〉 Torenia hybrida
〈223〉 Xaa(64) is Cys or Phe, Xaa(65) is Ser or Pro.
〈400〉 10
Met Val Asn Lys Arg His Ile Leu Leu Ala Thr Phe Pro Ala Gln Gly
1 5 10 15
His Ile Asn Pro Ser Leu Glu Phe Ala Lys Arg Leu Leu Asn Thr Gly
20 25 30
Tyr Val Asp Gln Val Thr Phe Phe Thr Ser Val Tyr Ala Leu Arg Arg
35 40 45
Met Arg Phe Glu Thr Asp Pro Ser Ser Arg Ile Asp Phe Val Ala Xaa
50 55 60
Xaa Asp Ser Tyr Asp Asp Gly Leu Lys Lys Gly Asp Asp Gly Lys Asn
65 70 75 80
Tyr Met Ser Glu Met Arg Lys Arg Gly Thr Lys Ala Leu Lys Asp Thr
85 90 95
Leu Ile Lys Leu Asn Asp Ala Ala Met Gly Ser Glu Cys Tyr Asn Arg
100 105 110
Val Ser Phe Val Val Tyr Ser His Leu Phe Ser Trp Ala Ala Glu Val
115 120 125
Ala Arg Glu Val Asp Val Pro Ser Ala Leu Leu Trp Ile Glu Pro Ala
130 135 140
Thr Val Phe Asp Val Tyr Tyr Phe Tyr Phe Asn Gly Tyr Ala Asp Asp
145 150 155 160
Ile Asp Ala Gly Ser Asp Gln Ile Gln Leu Pro Asn Leu Pro Gln Leu
165 170 175
Ser Lys Gln Asp Leu Pro Ser Phe Leu Leu Pro Ser Ser Pro Ala Arg
180 185 190
Phe Arg Thr Leu Met Lys Glu Lys Phe Asp Thr Leu Asp Lys Glu Pro
195 200 205
Lys Ala Lys Val Leu Ile Asn Thr Phe Asp Ala Leu Glu Thr Glu Gln
210 215 220
Leu Lys Ala Ile Asp Arg Tyr Glu Leu Ile Ser Ile Gly Pro Leu Ile
225 230 235 240
Pro Ser Ser Ile Phe Ser Asp Gly Asn Asp Pro Ser Ser Ser Asn Lys
245 250 255
Ser Tyr Gly Gly Asp Leu Phe Arg Lys Ala Asp Glu Thr Tyr Met Asp
260 265 270
Trp Leu Asn Ser Lys Pro Glu Ser Ser Val Val Tyr Val Ser Phe Gly
275 280 285
Ser Leu Leu Arg Leu Pro Lys Pro Gln Met Glu Glu Ile Ala Ile Gly
290 295 300
Leu Ser Asp Thr Lys Ser Pro Val Leu Trp Val Ile Arg Arg Asn Glu
305 310 315 320
Glu Gly Asp Glu Gln Glu Gln Ala Glu Glu Glu Glu Lys Leu Leu Ser
325 330 335
Phe Phe Asp Arg His Gly Thr Glu Arg Leu Gly Lys Ile Val Thr Trp
340 345 350
Cys Ser Gln Leu Asp Val Leu Thr His Lys Ser Val Gly Cys Phe Val
355 360 365
Thr His Cys Gly Trp Asn Ser Ala Ile Glu Ser Leu Ala Cys Gly Val
370 375 380
Pro Val Val Cys Phe Pro Gln Trp Phe Asp Gln Gly Thr Asn Ala Lys
385 390 395 400
Met Ile Glu Asp Val Trp Arg Ser Gly Val Arg Val Arg Val Asn Glu
405 410 415
Glu Gly Gly Val Val Asp Arg Arg Glu Ile Lys Arg Cys Val Ser Glu
420 425 430
Val Ile Lys Ser Arg Glu Leu Arg Glu Ser Ala Met Met Trp Lys Gly
435 440 445
Leu Ala Lys Glu Ala Met Asp Glu Glu Arg Gly Ser Ser Met Asn Asn
450 455 460
Leu Lys Asn Phe Ile Thr Arg Ile Ile Asn Glu Asn Ala Ser
465 470 475
〈210〉 11
〈211〉 335
〈212〉 PRT
〈213〉 Perilla frutescens
〈400〉 11
Met Ser Ser Ser Ser Ser Arg Arg Trp Arg Glu Asn Glu Gly Met Arg
1 5 10 15
Arg Thr Leu Leu Gly Leu Gly Leu Gly Gln Leu Val Ser Phe Asp Leu
20 25 30
Ala Ile Met Thr Phe Ser Ala Ser Leu Val Ser Thr Thr Val Asp Ala
35 40 45
Pro Leu Thr Met Ser Phe Thr Thr Tyr Thr Val Val Ala Leu Leu Tyr
50 55 60
Gly Thr Ile Leu Leu Tyr Arg Arg His Lys Phe Leu Val Pro Trp Tyr
65 70 75 80
Trp Tyr Ala Leu Leu Gly Phe Val Asp Val His Gly Asn Tyr Leu Val
85 90 95
Asn Lys Ala Phe Glu Leu Thr Ser Ile Thr Ser Val Ser Ile Leu Asp
100 105 110
Cys Trp Thr Ile Val Trp Ser Ile Ile Phe Thr Trp Met Phe Leu Gly
115 120 125
Thr Lys Tyr Ser Val Tyr Gln Phe Val Gly Ala Ala Ile Cys Val Gly
130 135 140
Gly Leu Leu Leu Val Leu Leu Ser Asp Ser Gly Val Thr Ala Ala Gly
145 150 155 160
Ser Asn Pro Leu Leu Gly Asp Phe Leu Val Ile Thr Gly Ser Ile Leu
165 170 175
Phe Thr Leu Ser Thr Val Gly Gln Glu Tyr Cys Val Lys Arg Lys Asp
180 185 190
Arg Ile Glu Val Val Ala Met Ile Gly Val Phe Gly Met Leu Ile Ser
195 200 205
Ala Thr Glu Ile Thr Val Leu Glu Arg Asn Ala Leu Ser Ser Met Gln
210 215 220
Trp Ser Thr Gly Leu Leu Ala Ala Tyr Val Val Tyr Ala Leu Ser Ser
225 230 235 240
Phe Leu Phe Cys Thr Leu Thr Pro Phe Leu Leu Lys Met Ser Gly Ala
245 250 255
Ala Phe Phe Asn Leu Ser Met Leu Thr Ser Asp Met Trp Ala Val Ala
260 265 270
Ile Arg Thr Phe Ile Tyr Asn Gln Glu Val Asp Trp Leu Tyr Tyr Leu
275 280 285
Ala Phe Cys Leu Val Val Val Gly Ile Phe Ile Tyr Thr Lys Thr Glu
290 295 300
Lys Asp Pro Asn Asn Thr Arg Ala Leu Glu Asn Gly Asn Leu Asp His
305 310 315 320
Glu Tyr Ser Leu Leu Glu Asp Gln Asp Asp Thr Pro Arg Lys Pro
325 330 335
〈210〉 12
〈211〉 468
〈212〉 PRT
〈213〉 Petunia hybrida
〈400〉 12
Met Val Gln Pro His Val Ile Leu Thr Thr Phe Pro Ala Gln Gly His
1 5 10 15
Ile Asn Pro Ala Leu Gln Phe Ala Lys Asn Leu Val Lys Met Gly Ile
20 25 30
Glu Val Thr Phe Ser Thr Ser Ile Tyr Ala Gln Ser Arg Met Asp Glu
35 40 45
Lys Ser Ile Leu Asn Ala Pro Lys Gly Leu Asn Phe Ile Pro Phe Ser
50 55 60
Asp Gly Phe Asp Glu Gly Phe Asp His Ser Lys Asp Pro Val Phe Tyr
65 70 75 80
Met Ser Gln Leu Arg Lys Cys Gly Ser Glu Thr Val Lys Lys Ile Ile
85 90 95
Leu Thr Cys Ser Glu Asn Gly Gln Pro Ile Thr Cys Leu Leu Tyr Ser
100 105 110
Ile Phe Leu Pro Trp Ala Ala Glu Val Ala Arg Glu Val His Ile Pro
115 120 125
Ser Ala Leu Leu Trp Ser Gln Pro Ala Thr Ile Leu Asp Ile Tyr Tyr
130 135 140
Phe Asn Phe His Gly Tyr Glu Lys Ala Met Ala Asn Glu Ser Asn Asp
145 150 155 160
Pro Asn Trp Ser Ile Gln Leu Pro Gly Leu Pro Leu Leu Glu Thr Arg
165 170 175
Asp Leu Pro Ser Phe Leu Leu Pro Tyr Gly Ala Lys Gly Ser Leu Arg
180 185 190
Val Ala Leu Pro Pro Phe Lys Glu Leu Ile Asp Thr Leu Asp Ala Glu
195 200 205
Thr Thr Pro Lys Ile Leu Val Asn Thr Phe Asp Glu Leu Glu Pro Glu
210 215 220
Ala Leu Asn Ala Ile Glu Gly Tyr Lys Phe Tyr Gly Ile Gly Pro Leu
225 230 235 240
Ile Pro Ser Ala Phe Leu Gly Gly Asn Asp Pro Leu Asp Ala Ser Phe
245 250 255
Gly Gly Asp Leu Phe Gln Asn Ser Asn Asp Tyr Met Glu Trp Leu Asn
260 265 270
Ser Lys Pro Asn Ser Ser Val Val Tyr Ile Ser Phe Gly Ser Leu Met
275 280 285
Asn Pro Ser Ile Ser Gln Met Glu Glu Ile Ser Lys Gly Leu Ile Asp
290 295 300
Ile Gly Arg Pro Phe Leu Trp Val Ile Lys Glu Asn Glu Lys Gly Lys
305 310 315 320
Glu Glu Glu Asn Lys Lys Leu Gly Cys Ile Glu Glu Leu Glu Lys Ile
325 330 335
Gly Lys Ile Val Pro Trp Cys Ser Gln Leu Glu Val Leu Lys His Pro
340 345 350
Ser Leu Gly Cys Phe Val Ser His Cys Gly Trp Asn Ser Ala Leu Glu
355 360 365
Ser Leu Ala Cys Gly Val Pro Val Val Ala Phe Pro Gln Trp Thr Asp
370 375 380
Gln Met Thr Asn Ala Lys Gln Val Glu Asp Val Trp Lys Ser Gly Val
385 390 395 400
Arg Val Arg Ile Asn Glu Asp Gly Val Val Glu Ser Glu Glu Ile Lys
405 410 415
Arg Cys Ile Glu Leu Val Met Asp Gly Gly Glu Lys Gly Glu Glu Leu
420 425 430
Arg Lys Asn Ala Lys Lys Trp Lys Glu Leu Ala Arg Glu Ala Val Lys
435 440 445
Glu Gly Gly Ser Ser His Lys Asn Leu Lys Ala Phe Ile Asp Asp Val
450 455 460
Ala Lys Gly Phe
465
Claims (11)
- 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자.
- 제1항에 있어서, 서열 7 내지 10 또는 12중 어느 하나에 기재된 아미노산 서열을 가지며 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질 또는 이의 아미노산 서열에 대하여 1개 또는 복수개의 아미노산의 부가, 결실 및/또는 다른 아미노산에 의한 치환에 따라 변형되는 아미노산 서열을 갖는 동시에 플라보노이드의 5위치에 배당체를 전이하는 활성을 유지하는 단백질을 암호화하는 유전자.
- 제1항에 있어서, 서열 7 내지 10 또는 12중 어느 하나에 기재된 아미노산 서열에 대하여 30% 이상의 상동성을 갖는 아미노산 서열을 가지며 또한 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자.
- 제1항에 있어서, 서열 7 내지 10 또는 12중 어느 하나에 기재된 아미노산 서열에 대하여 50% 이상의 상동성을 갖는 아미노산 서열을 가지며 또한 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자.
- 제1항에 있어서, 서열 7 내지 10 또는 12중 어느 하나에 기재된 아미노산 서열을 암호화하는 염기 서열의 일부 또는 전부에 대하여 5×SSC 및 50℃의 조건하에서 하이브리드화할 수 있으며 또한 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 암호화하는 유전자.
- 제1항 내지 제5항중 어느 한 항에 따른 유전자를 함유하는 벡터.
- 제6항에 따른 벡터에 의해 형질 전환된 숙주.
- 제1항 내지 제5항 중 어느 한 한 항에 따른 유전자에 의해 암호화되는 단백질.
- 제7항에 따른 숙주를 배양하거나 생육시키고 당해 숙주로부터 플라보노이드의 5위치에 배당체를 전이하는 활성을 갖는 단백질을 회수하는 것을 특징으로 하는 단백질의 제조방법.
- 제1항 내지 제5항중 어느 한 항에 따른 유전자가 도입되는 식물 또는 이와 동일한 성질을 갖는 이의 자손 또는 이들의 조직.
- 제10항에 따른 식물 또는 이와 동일한 성질을 갖는 이의 자손의 절화(切花).
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JP20057197 | 1997-07-25 | ||
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KR1019997002547A KR100718475B1 (ko) | 1997-07-25 | 1998-07-16 | 당전이 활성을 갖는 단백질을 암호화하는 유전자 |
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US (2) | US6855862B1 (ko) |
EP (1) | EP0967283B1 (ko) |
JP (1) | JP4293641B2 (ko) |
KR (1) | KR100718475B1 (ko) |
CN (1) | CN1322130C (ko) |
AT (1) | ATE387493T1 (ko) |
AU (1) | AU754464B2 (ko) |
CA (1) | CA2266421C (ko) |
DE (1) | DE69839178T2 (ko) |
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KR20170051138A (ko) * | 2015-10-30 | 2017-05-11 | 고려대학교 산학협력단 | 단순 페놀릭 배당 유도체, 이의 제조방법 및 이를 함유하는 피부미백 또는 주름개선용 조성물 |
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NZ507563A (en) * | 1999-02-16 | 2004-02-27 | Suntory Ltd | Gene encoding a protein having activity of transferring sugar onto aurone |
GB0002814D0 (en) | 2000-02-09 | 2000-03-29 | Univ York | Nucleic acids and their uses |
JP4827043B2 (ja) | 2000-06-02 | 2011-11-30 | サントリーホールディングス株式会社 | 新規脂肪族アシル基転移酵素遺伝子 |
EP1291418B1 (en) * | 2000-06-02 | 2009-05-06 | International Flower Developments Proprietary Limited | Novel glycosyltransferase genes |
ATE525465T1 (de) * | 2005-10-20 | 2011-10-15 | Local Independent Administrative Institution Aomori Prefectural Ind Technology Res Ct | Neue gene für transferasen aromatischer acylgruppen |
JP2010532385A (ja) * | 2007-07-02 | 2010-10-07 | ユ,ミン | 複合的癌治療の方法、組成物および標的 |
JP5553317B2 (ja) * | 2009-04-24 | 2014-07-16 | サントリーホールディングス株式会社 | 花弁で機能するシソ由来のプロモーター |
EP2664671A4 (en) | 2011-01-14 | 2014-09-10 | Suntory Holdings Ltd | New Glycosyltransferase and its use |
JP6147195B2 (ja) | 2012-01-17 | 2017-06-14 | サントリーホールディングス株式会社 | 新規糖転移酵素遺伝子及びその使用 |
CN103348799A (zh) * | 2013-07-19 | 2013-10-16 | 镇江瑞繁农艺有限公司 | 美女樱多次采种栽培方法 |
EP3114210A2 (en) * | 2014-03-07 | 2017-01-11 | Evolva SA | Methods for recombinant production of saffron compounds |
CN103952414A (zh) * | 2014-05-22 | 2014-07-30 | 中南民族大学 | 一种烟草糖基转移酶诱导型启动子及其应用 |
CN109868265B (zh) * | 2017-12-01 | 2023-04-25 | 中国科学院分子植物科学卓越创新中心 | 新型糖基转移酶及其应用 |
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DK0656940T3 (da) | 1992-07-30 | 2004-02-09 | Int Flower Dev Pty Ltd | Gensekvenser, som koder for glykosyltransferase-enzymer, og anvendelser deraf |
JP2987277B2 (ja) * | 1993-08-31 | 1999-12-06 | 日新製鋼株式会社 | 金属溶解炉の水冷構造 |
US6489541B1 (en) * | 1994-06-24 | 2002-12-03 | Michigan State University Board Of Trustees | Genetic control of plant hormone levels and plant growth |
DE69603609T2 (de) | 1995-06-14 | 2000-03-23 | Exxon Chemical Patents, Inc. | Verbesserte elastomerische strangpressprofile |
EP0771878A1 (en) * | 1995-10-31 | 1997-05-07 | Plant Genetic Systems N.V. | Plants with reduced glucosinolate content |
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1998
- 1998-07-16 AU AU82432/98A patent/AU754464B2/en not_active Ceased
- 1998-07-16 AT AT98932550T patent/ATE387493T1/de not_active IP Right Cessation
- 1998-07-16 CN CNB988012146A patent/CN1322130C/zh not_active Expired - Fee Related
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- 1998-07-16 JP JP50964799A patent/JP4293641B2/ja not_active Expired - Fee Related
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- 1998-07-16 KR KR1019997002547A patent/KR100718475B1/ko not_active IP Right Cessation
- 1998-07-16 WO PCT/JP1998/003199 patent/WO1999005287A1/ja active IP Right Grant
- 1998-07-16 DE DE69839178T patent/DE69839178T2/de not_active Expired - Lifetime
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Cited By (1)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
KR20170051138A (ko) * | 2015-10-30 | 2017-05-11 | 고려대학교 산학협력단 | 단순 페놀릭 배당 유도체, 이의 제조방법 및 이를 함유하는 피부미백 또는 주름개선용 조성물 |
Also Published As
Publication number | Publication date |
---|---|
ES2299210T3 (es) | 2008-05-16 |
EP0967283A1 (en) | 1999-12-29 |
JP4293641B2 (ja) | 2009-07-08 |
DE69839178T2 (de) | 2009-02-19 |
AU8243298A (en) | 1999-02-16 |
CA2266421A1 (en) | 1999-02-04 |
NZ335001A (en) | 2001-03-30 |
KR100718475B1 (ko) | 2007-05-16 |
US20050257291A1 (en) | 2005-11-17 |
EP0967283A4 (en) | 2004-06-09 |
ATE387493T1 (de) | 2008-03-15 |
CA2266421C (en) | 2011-02-15 |
CN1237206A (zh) | 1999-12-01 |
DE69839178D1 (de) | 2008-04-10 |
EP0967283B1 (en) | 2008-02-27 |
CN1322130C (zh) | 2007-06-20 |
US6855862B1 (en) | 2005-02-15 |
AU754464B2 (en) | 2002-11-14 |
WO1999005287A1 (fr) | 1999-02-04 |
US7501271B2 (en) | 2009-03-10 |
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