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“Acapulco” 오리엔탈 나리에서 분리한 chalcone synthase을 인지하는 약 특이 프로모터의 구명

The characterization of anther-specific promoter encoding chalcone synthase isolated from Lilium Oriental hybrid cv. Acapulco

Korean Journal of Breeding Science 2017;49(3):190-199.
Published online: August 31, 2017

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*Corresponding Author (seji00@korea.kr, +82-63-238-4660, +82-63-238-4654)
• Received: June 27, 2017   • Accepted: August 2, 2017

© Korean Society of Breeding Science. All rights reserved.

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  • Anthocyanins are the major pigments contributing to flower coloration. A 1584 bp 5’ upstream sequence of ALCHS2 gene was isolated from Acapulco lily (Lilium Oriental hybrid cv. Acapulco). Computer-based analyses (GeneScan, AtPAN) predicted a CAATBOX1 and putative transcription factor-binding sites, including tissue-specific elements. When gALCHS7 promoter–gus fusion was introduced to petunia (‘Dream Red’), all ten putative transgenic plants showed localized GUS activity in the anther, but five of them also showed weak GUS activity in the ovule. No distinctive signal in the leaf and petal was detected in the same stage. To clearly determine the operation of the promotor region, anther and ovule tissues of transgenic line 6 were fixed in paraffin for dark-field analysis. At 1 cm length of floral bud, a GUS signal was not observed in the anther, but weak expression was observed in the ovule. Before anthesis, GUS protein was highly expressed in the pollen, endothecium, and epidermis. Fluorometric GUS assays of individual organs taken from four transgenic plants demonstrated that all lines showed high GUS activity in the anther compared to 35S CaMV promoter (pBI1 121), except line 34. Using the truncated promoters by cis-acting elements, we found that minimal region (gALCHS7-7, 270 bp) displayed GUS expression only in the anther, though at weaker activity than in the original promoter.
화색은 화훼작물에서 소비자의 주의를 끌어 판매를 유도할 수 있는 매우 중요한 특성이다. 플라보노이드 합성경로에서 다양 한 구조 및 조절유전자들이 확인되었는데 이중 chalcone synthase (CHS, EC2.3.174)는 가장 널리 연구된 유전자 중 하나이다(Liu et al. 2011). CHS는 플라보노이드 생합성에서 첫번째 반응을 촉매하는데(koes et al. 1989) malonyl-coenzyme A (CoA)의 3개의 aetate를 응축하여 4-coumaroyl-CoA와 함께 naringenin chalcone을 형성한다(Heller & Hahbrock 1980, Martin 1993). 현재까지 약 650개의 CHS 및 CHS유사 유전자가 식물의 단자엽과 쌍자엽을 통틀어 보고되었다(Yang & Gu 2006). CHS 유전자들의 발현은 대부분이 꽃과 식물의 발달과정 에서 발현된다고 밝혀져 있지만(Nakatsuka et al. 2003, Farzad et al. 2003, Han et al. 2005) 다양한 식물에서 광이나(Farzad et al. 2003, Meng et al. 2004) 상처 혹은 뿌리에 특이적으로도 발현하는것으로 알려져 있다(McKhann & Hirsch 1994, Ito et al. 1997, Yi et al. 2010). 화색이 중요한 화훼작물중에서는 페튜니아에서 최소 8개 이상(Koes et al. 1989), 나팔꽃에서는 5개의 CHS유전자가 보고 되었으며(Yang et al. 2004) Lillium에 서는 세 개의 CHS에 대해서 구명된 바 있다(Nakatsuka et al. 2009). 최근에 거베라에서 확인된 3개의 CHS유전자들은 특히 약의 sporopollenin생합성과 exine의 형성에 관여한다고 한다 (Kontturi et al. 2017). 모든 작물에 해당하지는 않으나 실제로 몇몇 작물에서 플라보노이드가 화분의 발아와 임성에 중요한 역할을 한다고 알려져 있는데(Coe et al. 1981, Taylor & Jorgensen. 1992, van derMeer et al. 1992, Fischer et al. 1997, Napoli et al. 1999) 배추(Shen & Hsu 1992), 벼(Hihara et al. 1996), 담배(Atanassov et al. 1998) 및 달맞이장구채 (Barbacar et al. 1997)에서 분리한 CHS 염기서열에 유사한 유전자들이 약의 tapetum세포에서 발현이 확인되었다. 약 특이 적 CHS 유사 효소들은 아미노산, 염기서열 및 촉매반응은 일반 적인 CHS와 유사하지만 기질에 대한 선호도가 다른것으로 확인 되어서 종종 anther-specific CHS-like enzyme (ASCLs)라고 구별하여 언급하기도 했다(Kontturi et al. 2017). CHS 유전자의 기능처럼 CHS를 인지하는 프로모터도 여러 시그널에 반응하는 것으로 확인되었다. 페튜니아에서 분리한 CHS 인지 프로모터 (220 bp)의 경우 꽃 특이적 발현을 보였고(van der Meer et al. 1990) 포플라에서 분리한 프로모터의 경우 환경적인 요인(상 처 등)에 반응하여 발현되었으며(Sun et al. 2011) 파슬리의 CHS프로모터는 빛에 반응하여 작동하였다(Weisshaar et al. 1992). 페튜니아에서 분리한 프로모터인 chiA와 chiB는 약특이 (화분포함)발현을 보였고 담배류의 화분에서 발현되는 CHS유 전자의 프로모터는 발달단계상 약의 소포자 단계에서 확인되었 다(van Tunen et al. 1990, Atanassov et al. 1998).
나리는 독특한 화형의 꽃과 다양한 화색 때문에 가장 중요한 화훼작물의 하나로 간주된다. 지금까지 밝혀진 나리 속은 90종이 상으로 표현형에 따라 7개의 그룹으로 분류할 만큼 다양한 형태 와 품종이 개발되어 있다(The Royal Horticultural Society 2014). 나리가 다양한 색과 형태 및 품종을 지니고 있긴 하지만 절화 수명이 짦고, 보라색계열의 화색이 없으며, 특히 오리엔탈 나리의 경우 개화 후 성숙한 약이 터지면서 꽃잎에 얼룩이 생겨 상품성을 떨어뜨린다. 일반적으로 나리의 경우 형질전환이 매우 어려운 것으로 알려져 있지만 최근에 아카풀코를 비롯한 나리의 형질전환 사례가 보고가 되고 있어서 육종으로 도입이 어려운 형질의 유전자를 이용한 형질전환이 불가능한 것만은 아니다 (Azadi et al. 2004, Hoshi et al. 2004). 본 실험에서는 오리엔탈 나리인 아카풀코에서 약 특이적 프로모터를 개발하여 약에서의 유전자 발현을 연구하는데 이용하고 농업적으로 나리의 상품성 을 높이는데 활용하고자 한다.
식물재료 및 프로모터 분리
본 실험에 사용한 아카풀코 나리(Lilium Oriental hybrid cv. Acapulco)는 Netherland Bulb Company (USA, Easton Pennsylvania)에서 구매하였으며 구의 직경이 16-18cm 되는 것을 피트모스와 펄라이트를 섞은 상토에 심어 국립원예특작과 학원 온실에서 15-25°C로 재배하였다. 잎으로부터 Murray 와 Thompson (1980)의 방법에 따라서 분리한 게놈 DNA를 가지고 Lambda Fix Ⅱ/XhoⅠ kit (Startagene)와 ZAP-cDNA Gigapack cloning kit (Stratagene)을 사용하여 제조사의 지침에 따라서 라이브러리를 제작하였다. 라이브러리 screenig에 사용 한 탐침은 꽃잎과 약에서만 발현이 되는 ALCHS2 cDNA (GenBank accession no. AF169799)를 RediprimeTM Ⅱ DNA Labeling system (Amersham)을 이용하여 제조사의 지침에 따 라 [α-32P] dCTP로 표지하여 사용하였다. 분리한 gALCHS7 (Genome Acapulco Lily Chalcone Synthase 7: 약 15.0 kbp)를 EcoRⅠ으로 잘라서 ALCHS2를 포함하는 6.2 kbp의 클론을 확보하였다.
CHS 게놈 유전자의 염기서열 분석 및 전사조절 부위조사
약 6.0 kbp의 gALCHS7 클론(GenBank accession no. KY883978)의 염기서열을 분석하기 위해서 몇 개의 제한효소로 절단하여 제한효소 지도를 작성하였다. 이 지도에 따라서 SalⅠ 과 EcoRⅠ으로 절단하여 pBluescript Ⅱ SK (+) 벡터로 subcloning을 수행한 다음, ABI Prime 310 Genetic Analyzer (PE Applied Biosystems)로 분석하여 완전한 전체 염기서열을 얻었다. 얻어진 6203 bp의 염기서열은 genescan (http://genes.mit.edu)을 이용하여 exon과 intron 부위를 확인하였고 AtPAN (http:// atpan.itps.ncku.edu.tw/index.php), Chen et al. 2012) 에 전송하여 전사조절 부위를 분석하였다.
벡터제작 및 형질전환
6.2 kbp의 게놈클론에서 5'-flanking 부위(gALCHS7) 중 개시 코돈앞의 5' UTR부분(1584 bp)을 PCR을 이용하여 증폭한 다음 벡터 제작에 사용하였다. 증폭에 사용한 primer는 gSalⅠ(Upper primer : 5'-TAGCACATGTGCAGGG TCG ACA-3') 과 gXba Ⅰ(Lower primer : 5'-ATTTCTAGAACCGAAGATACTA-3') 으로 증폭 후 SalⅠ과 EcoRⅠ로 절단한 다음, 분리하여 동일한 제한효소로 절단한 벡터pBI101(Jefferson et al. 1987)과 연결하 여 gALCHS7 promoter::uidA벡터를 제작하였다. 약에서만 작동 하는 부위를 찾기 위해 작성한 gALCHS7-7 promoter::uidA의 벡터는 프로모터 염기 서열 –266에서 gSalⅠ-1(Upper primer : 5' CACTCAGGGGACTTGCTACGAACCAAA-3')와 gXbaⅠ 프라이머를 이용하여 동일한 방법으로 벡터를 제작하였다 (Fig. 1). 페튜니아 (Dreams Red, PanAmerican Seed)로의 형질전환은 Agrobacterium stain C58C1을 이용하여 Horsh 등 (1985)의 방법을 변형한 leaf disk법을 사용하였다. 형질전환체 선발은 kanamycin (50 mg·L-1)이 포함된 배지에서 4주간 키운 shoot를 다시 kanamycin (100 mg·L-1) 포함 배지로 옮겨 3주간 배양하여 선발하였다. 선발 형질전환체는 kanamycin이 포함 되지 않은 발근 배지로 옮겨 약 4-6주간 발근을 유도한 뒤 온실로 옮겨 순화시켰다. 선발한 형질 전환체의 잎으로부터 DNeasy Plant Mini Kit (Qiagen, Germany)을 이용하여 게놈 DNA를 분리한 다음 NPT Ⅱ primer (Primer 1: 5'- GAG GCT ATT CGG CTA TGA CTG –3', Primer 2: 5' -ATC GGG AGC GGC GAT ACC GTA –3')로 증폭하여 형질 전환여부를 확인하였다.
Fig. 1
Nucleotide sequence from lily genomic clone gALCHS7. Only 1584 nt of the genomic sequence are shown. The start codon of ALCHS2 are shown in upper case (M). Computer deduced putative cis-elements are boxed (http://atpan.itps.ncku.edu.tw/index.php). Putative anther specific promoter region (gALCHS7-7) was amplified with forward primer (underlined) and the same reverse primer of gALCHS7 promoter.
KJBS-49-190_F1.gif
Histochemical 분석
GUS 활성은 페튜니아 형질 전환체의 각 기관을 이용하여 조사하였다. 페튜니아 꽃에서 개약전 단계의 잎과 꽃잎, 약, 암술, 자방을 면도칼로 자른 다음 12시간동안 37℃에서 GUS용액 (De Block M & Debrouwer D 1992)에 침지하였다. 이때 형질 전환이 되지 않은 조직에서 내생 GUS의 유사발현을 억제하기 위해서 20% methanol을 첨가하여 사용하였다 (Kosugi et al. 1990). 기관별 GUS단백질 활성비교는 페튜니아 형질 전환체의 약과 자방을 이용하여 조사하였다. 페튜니아 꽃에서 개약전 단계의 약, 자방을 각각 채취하여 Jefferson 등(1987)의 방법에 따라서 각 기관별로 단백질을 추출하였다. 추출한 단백질은 spectrophotometer (Bio-Rad)로 정량한 다음 Jefferson 등 (1987)의 방법에 따라서 fluorimeter (Bio-Rad)를 이용하여 비 교하였다. 이때 사용한 대조구로는 비형질전환 식물체의 각 기관 단백질과 35S CaMV (Cauliflower Mosaic Virus)프로모터에 GUS가 연결되어 있는 pBI121을 형질전환한 식물체의 각 기관 으로부터 분리한 단백질을 이용하였다. 측정된 수치들은 SAS 프로그램(Hussein et al. 2000)으로 유의차를 분석하였다.
Dark-field기법을 이용한 GUS 발현 분석
약 기관내에서 GUS발현을 자세히 관찰하기 위해서 다음 두 가지 방법을 조합하여 시료를 고정한 다음 dark-field로 관찰하였 다 (Ausubel et al. 1995, Bossinger et al. 1996). GUS활성을 보인 조직을 GUS염색한 후 상온에서 다음과 같은 용액(50% 에탄올, 5% 아세틱에시드, 3.7% 포름알데하이드)에 12시간 침 지하여 고정하였다. 에탄올 시리즈(70%, 85%, 95%)를 이용하 여 탈수 후 크실렌(xylene)과 에탄올 시리즈(1:3, 1:1, 3:1)을 이용하여 각 1시간씩 침지시킨 다음 100% 크실렌으로 각 1시간 씩 2회 더 갈아준다. 약 5-10 ml의 크실렌을 넣은 후 동일한 양의 파라핀(paraplast, Fisher)을 넣고 60℃오븐에서 12시간 둔다. 잘 녹은 파라핀은 버리고 새로운 파라핀을 붓는 과정을 2-3회 반복한 다음 틀에 부어 고정한 후 절단에 알맞은 모양으로 다듬어 둔다. 파라핀절단기기(Leica)를 이용하여 10-15μm로 절단하여 poly-prep slide (sigma)에 고정한 다음 37℃에 12시간 정도 말린다. 현미경관찰을 위해서 시료가 고정된 슬라이드를 100% 크실렌에 12시간 녹인다음 크실렌(xylene)과 에탄올 시리 즈(3:1, 1:1, 1:3)을 이용하여 탈수한 후 Permount (Fisher)를 도포하여 커버슬립을 덮어 고정하고 말린 후 dark-field용 필터가 장착된 현미경(Leica)에서 관찰하였다.
게놈클론 gALCHS7의 분리와 염기서열 분석
오리엔탈 나리 아카풀코(Lilium Oriental hybrid cv. Acapulco) 의 게놈 라이브러리는 Lambda Fix Ⅱ/XhoⅠ 벡터를 이용해서 만들었고 ALCHS2 cDNA (AF169799)를 탐침으로 게놈 클론인 gALCHS7 클론을 분리했다. 잎의 게놈 DNA로 만들어진 게놈 라이브러리는 5 × 105 pfu였으며 평균 insert의 크기는 9-23 kbp였다. 꽃잎과 약에서 발현을 보이는 ALCHS2의 3' end 부분 을 탐침으로 하여서 30만개의 게놈유전자를 조사한 결과 18개 의 후보클론을 확보할 수 있었다(data not shown). 선발한 gALCHS7은 약 6.0 kbp의 게놈 클론이 cDNA의 ORF부분을 포함하고 있는 것을 서던블럿으로 확인한 다음 pBluescript Ⅱ SK (+) 벡터로 옮긴 뒤 전체 염기서열을 얻었다 (6203 bp). 전체 염기서열 분석결과 탐침으로 사용한 ALCHS2 cDNA의 ORF부 분과 gALCHS7의 exon 부분의 아미노산 서열을 비교한 결과 96.8% 일치를 보였다 (data not shown). 이러한 결과로 볼 때 gALCHS7 게놈 클론은 CHS유전자의 프로모터 부분을 포함하고 있을 것으로 생각되었다.
gALCHS7 프로모터의 전사조절 부위조사
gALCHS7 게놈클론에서 5' UTR부위의 염기서열을 AtPAN (http://atpan.itps.ncku.edu.tw/index.php) 에 전송하여 cis-acting elements를 분석하였다(Fig. 1). 진핵생물에 일반적으로 보고된 연 속적 염기서열(consensus sequence)외에 LTRECOREATCOR15 염기서열은 애기장대에서 저온에 반응하는 인자로 알려져 있으며 ASF-2는 GATABOX로 불리는 인자로 식물형질전환에 많이 사용하는 CaMV 35S 프로모터에서 보고된 GATA모티브이다. 이 모티브는 페튜니아 chlprophyll a/B binding protein인 Cab22 의 프로모터에서 발견되었는데 발현양을 증가시키거나 빛 혹은 조직특이적 발현에 관여한다고 한다. MYB2는 페닐알라닌 계열 의 이차 대사산물인 리그닌, 플라보놀, 안토시아닌의 합성에 관여하는 유전자의 프로모터부분에서 c-Myb과 P-box를 인지하 는 부위로 완두에서 보고된 바 있는 전사인자이다(Uimari & Strommer 1997). GTGANTG10 (GTGA)은 담배의 화분발현 유전자g10에서 발현하는 pectate lyase의 프로모터에서 보고된 인자로 gALCHS7 프로모터에서는 두 개의 인자가 존재하는 것으 로 확인되었다. GTGANTG10인자는 토마토의 LAT52 유전자 의 화분특이적 발현에 영향을 주는 인자로 보고된 바 있는 POLLEN1LELAT52인자와 더블어 gALCHS7 프로모터의 약 에서의 발현에 영향을 주는 주요한 인자로 생각된다. 두 개가 확인된 OSE2ROOTNODULE인자의 경우 기관특이적인 발현 에 영향을 주는 연속적인 염기서열로 보이며 C1의 경우 옥수수의 안토시아닌 합성을 조절하는 조절인자중의 하나로 보고된 바 있다(Bodeau JP & Walbot 1996). LCR1은 PCP (pollen coat protein) 패밀리 유전자의 염기서열과 유사한 시스테인 리치 단백질 패밀리를 인지하는 것으로 알려져 있다(Lin et al. 1999). gALCHS7 프로모터는 개시시작부위에 결합 일반적으로 프로모 터에서 보고되는 CAATBOX1와 TATABOX5는 유전자 발현 개시 앞에서 확인되나 TATABOX5의 경우 유전자발현 시작점 에서 상당히 떨어진 지점(-603)에서 나타났다.
gALCHS7 프로모터의 페튜니아 형질전환 및 GUS발현 분석
gALCHS7 promoter::uidA 벡터를 페튜니아에 leaf disk (Horsh et al. 1985)방법을 이용하여 형질전환을 수행하였다. 나리에서 분리한 유전자는 나리에서 발현을 확인하는 것이 맞으 나 나리의 형질전환은 아직 일반적이지 않고 gALCHS7 프로모터 가 암호화한 CHS 유전자는 페튜니아에서 12개의 이상의 유전자 가 존재하고 이중 4개가 발현된다고 보고 되어 있어서(Koes et al. 1989) 프로모터를 작동시키기 위한 요소들이 충분할 것으 로 판단되어 형질전환에 이용하였다. 형질전환 후 선발한 신초를 발근시켜 온실에서 순화시킨 형질전환체 개체에서 개약전 단계 의 잎, 약, 암술(암술머리, 암술대), 자방(배주포함), 꽃잎을 분리 하여 각각 GUS 용액에 침지하여 활성을 측정하였다. 10개의 개체 모두가 약에서 강한 GUS 활성을 보였으나 일부 개체에 서 단면으로 절단한 자방에서 약한 GUS발현이 확인되었다 (Fig. 2). 정확히 약의 어느 부위에서 프로모터가 작동하는지 확인하기 위해 페튜니아 형질전환체를 크게 두 단계의 약과 자방(꽃봉오리크기 1cm, 개약 전)을 paraffin에 고정하여 dark-field기법를 이용하여 관찰하였다(Fig. 3). 관찰결과 자방 과 약에서 동시에 발현을 보인 6번개체의 경우 tapetum조직이 무너지면서 화분이 발달하는 1cm 꽃봉오리의 약(Fig. 3F)에서 는 발현을 확인 할 수 없었지만 동일한 단계의 자방에서는 약한 GUS발 현을 확인할 수 있었다(Fig. 3I). 개약전 단계에서는 (Fig. 3G) 약의 endothecium, epidermis에서 강한 GUS발현이 나타났으며 화 분에서도 전체적으로 강한 시그널을 보였고 GUS염색과 동일하 게 자방에서 GUS발현이 있었다. 약에서만 발현이 확인된 18번 개체의 경우에는 현미경을 통해서도 약에서만 발현되며 두 단계 의 성숙한 자방에서는 GUS 발현을 확인 할 수 없었다(data not shown).
Fig. 2
Histochemical localization of GUS activity transgenic petunia plants transformed with gALCHS7::uidA. A, non-transgenic plant; B, transgenic petunia harboring the gALCHS::uidA gene; C, transgenic petunia harboring the pBI121. An, anther ; Ov, ovule ; Pe, petal.
KJBS-49-190_F2.gif
Fig. 3
Dark-field analysis of a non-transgenic (A-E) and gALCHS7::uidA transformed plants(#6) (F-J). GUS expression at bud stage of 1 cm in size (A and F), before anthesis (B and G), and mature pollen (C and H). Longitudinal sections of ovule at bud stage of 1 cm in size (D and I), and before anthesis (E and J). Scale Bars, 1cm.
KJBS-49-190_F3.gif
gALCHS7 프로모터의 활성분석
프로모터의 활성을 측정하기 위해서 각 기관별로 단백질을 Jefferson 등의 (1987)방법에 따라서 단백질을 추출한 다음 정량 하고 flurometer (Bio-Rad)를 이용하여 GUS활성을 측정하였 다. 측정한 결과는 SAS program (Hussein et al. 2000)을 이용하 여 통계분석을 수행하였고 비교 대조구로는 페튜니아 비형질 전환체와 35S CaMV프로모터가 GUS를 발현시키는 pBI121을 사용하였다. GUS염색결과와 마찬가자로 6번 개체의 약이 35S 프로모터와 비교해서 약 3배정도 높은 활성을 나타냈다(Fig. 4). 34번을 제외한 나머지 개체들도 약에서 최소 1.5배 이상의 GUS 활성을 보였다. Van Tunen 등(1990)의 보고에 따르면 페튜니아 의 암술과 자방이 내재적으로 상당히 높은 GUS활성을 보인다는 보고가 있는데 본 실험의 대조구에서는 형질전환체에 비해 높은 활성을 보이지는 않았다. pBI121 벡터가 삽입되어 있는 형질전 환체의 자방과 거의 같은 수준으로 gALCHS7 포함 형질전환체의 자방도 GUS발현을 보였는데 37번 형질전환체의 경우 6번 개체 와 마찬가지로 약에서도 발현하지만 자방에서도 GUS발현이 높은 개체인 것으로 확인되었다.
Fig. 4
Changes in GUS activity drived by gALCHS7::uidA promoter and pBI 121 (35S CaMV::uidA) in petunia anther and ovules. GUS activity is averaged from three independent experiments. Vertical bars indicates standard deviation. The letters above colums indicate statistically significant differences at P<0.05 by SAS software (version SAS 8.2).
KJBS-49-190_F4.gif
약 특이 발현 프로모터(gALCHS7-7)의 분리 및 GUS발현 확인
gALCHS7프로모터의 경우 전형질전환체의 약에서 발현이 되 었으나 일부의 형질전환체에서는 자방에서도 작동하는 것으로 확인되었다. 약에서만 작동하는 부위를 찾기 위해 프로모터를 전사조절부위에 따라 약 100-200 bp절단하여 각각 벡터를 제작 한 다음 형질전환하여 GUS발현을 확인하였다(data not shown). 실험결과 -266에서부터 개시코돈 전까지의 프로모터부위 (gALCHS7-7)가 삽입된 페튜니아 형질전환체의 약에서 GUS발 현을 확인하였다(Fig. 5). 각 기관별로 GUS염색을 수행한 결과 약에서만 GUS가 발현되었고 다른 기관에서는 전혀 GUS발현이 확인 되지 않아서 이 부위가 처음에 분리한 1584 bp의 프로모터 에서 약 특이 발현을 주도하는 부위인것으로 확인할 수 있었다. gALCHS7 프로모터와 마찬가지로 dark-field기법을 이용하여 약에서의 발현 부위를 확인한 결과 gALCHS7 프로모터에 비해 활성이 좀 떨어지는 것으로 보이지만 발현부위는 개약 직전 약의 지지조직(endothecium, epidermis)과 화분에서만 발현이 되는 것을 볼 수 있었다(Fig. 6).
Fig. 5
Histochemical localization of GUS activity in petunia plants under gALCHS7-7 promoter. A, non-transgenic plant; B-C, transgenic petunia harboring the gALCHS7-7::uidA gene (B, line 16 and C, line 21). An, anther ; Ov, ovule ; Pe, petal
KJBS-49-190_F5.gif
Fig. 6
Representative results of gALCHS7-7 promoter through dark-field analysis. Stained anthers from Fig. 5 were embedded in paraffin and sectioned. A-B; anther from non-transgenic plants, C-F; transgenic petunia harboring the gALCHS7-7::uidA gene (C-D, line 16 and E-F, line 21). Scale Bars, 1cm.
KJBS-49-190_F6.gif
약은 꽃의 주요한 기관으로 여러 개의 조직으로 구성되었으며 생식에 중요한 역할을 하는 화분을 형성하여 수정을 위해 방출하 는 과정에 복합적인 유전자군들이 관여한다고 알려져 있으나 이중 아주 일부만이 약 특이적인 유전자일 것으로 추정된다(Liu et al. 2013). 나리의 경우 게놈 크기가 36GB로 게놈 사이즈가 상당히 큰 작물 중 하나로, 최근의 보고에 따르면 게놈의 크기가 큰 식물체는 유전자의 삽입, 반복 등이 빈번하여 게놈크기가 커진 것이라는 보고가 있는 것으로 볼 때 (Heslop-Harrison 1997), 나리게놈에도 이러한 transposon이 다수 존재할 것으로 추측된다. 실제로 처음에 분리한 2개의 클론에서 나머지 하나는 염기서열 분석결과 5' flanking부위에 transposon삽입이 확인되 어서 사용불가한 것으로 확인되었으며 실제 나리게놈에는 이러 한 transposon이 다수 존재할 것으로 추정된다. 나리에서 분리한 프로모터이므로 나리에서 발현시키는 것이 가장 적절할 것이나 나리의 경우 형질전환이 쉽지 않아 화색유전자가 많이 알려진 페튜니아를 이용하여 프로모터의 발현을 확인하였다. 페튜니아 의 게놈에는 12개의 CHS 유전자가 있으며 이중 4개의 유전자만 이 발현된다고 보고된 바 있다 (Koes et al. 1989). 나리에서 분리한 CHS 프로모터가 페튜니아에서 기관특이적으로 작동할 수 있었던 것은 gALCHS7이 가지고 있는 cis-acting element 외에도 페튜니아가 지닌 CHS 유전자 발현에 작용하는 trans-acting element의 조건도 충분했기 때문에 기관특이적인 발현을 보일 수 있었던 것으로 생각된다. 페튜니아로의 형질전환 후 GUS염색과정을 통해 기관발현을 확인할 수 있었지만 약의 어느 부위에서 정확히 발현되는지를 확인하기 위해서 꽃봉오리 크기가 1cm정도 일 때와 개약직전의 약을 고정하여 dark-field기 법으로 관찰하였다. 꽃봉오리가 1cm인 단계는 페튜니아 발달상 B1에 해당하는 시기로 자방은 성숙하여서 integument를 형성하 는 시기이며 화분은 uninucleate 소포자들이 각각 분리하여 화분 낭에서 자유로이 떨어져 있는 상태라고 알려져 있다(Decroocq- Ferrant et al. 1995). 실제 파라핀 고정 후 절단하여 관찰한 결과 화분이 tapetum에서 분리되어 화분낭으로부터 분리되고 있었으며 개약전의 약은 이미 성숙한 화분으로 가득 차 있었다. Dark-field기법을 이용한 관찰에서 1cm단계의 약에서는 발현을 확인할 수 없었으나 개약전 약에서는 강한 발현이 확인 되었다. 자방의 경우 1cm 단계부터 약한 발현을 보였는데 전사인자를 조사한 결과에서는 특별히 자방의 발현에 관련된 전사인자는 확인 할 수 없었는데 아마도 꽃에 관련되어 있는 몇 개의 전사인자 들이 관여할 것으로 추정된다. 유전자 분리에 사용한 오리엔탈 나리인 아카풀코의 꽃에서 약은 꽃봉오리가 열리기 전에는 노란 색의 약주머니를 띄고 있다가 성숙하면서 적벽돌색을 띠면서 개약시 동일한 색의 화분을 터트린다. 안토시아닌 합성경로에서 CHS유전자는 플라보노이드 합성의 초반에 관여하는 것으로 알려져 있는데 오리엔탈 나리인 아카풀코의 약은 화분이 발달하 는 약의 후반기 생장에서 적벽돌색을 만들어 내는 펠라고니딘을 합성할 것으로 추정된다(Tanaka et al. 2008). 따라서 CHS를 인지하고 있는 gALCHS7프로모터는 이 과정에서 작동하여 약 특이적 발현을 유도할 것으로 추정된다. 처음에 분리한 1584bp 프로모터(gALCHS7)가 삽입된 형질전환체가 약에서 발현이 강 했지만 일부 개체에서 자방에서의 발현이 확인되어서 약 특이적 인 발현을 주도하는 부위를 찾기위해 cis-elements에 따라 약 150-200bp씩 절단하여 6개의 벡터를 제작하였다(data not shown). 5'-upstream 절단 백터중 gALCHS7-7 앞부분의 프로모 터들은 gALCHS7프로모터와 마찬가지로 모두 약에서 발현은 강하게 나왔지만 일부 개체들의 자방에서 여전히 발현이 확인되 었다. 그러나 gALCHS7-7 promoter::uidA 벡터의 발현 강도가 약간 줄어들긴 했지만 유일하게 약에서만 발현되었다. gALCHS7 전반에 걸쳐 담배의 화분 발현 유전자에서 확인된 GTGANTG10 (GTGA)이나 POLLEN1LELAT52와 같은 토 마토의 화분 발현 인자을 인지하는 모티브 등이 확인 되었는데 gALCHS7-7프로모터의 경우 PCP (pollen coat protein)의 염기 서열과 유사한 시스테인 리치 단백질을 인지하는 부분을 포함하 고 있으며 개약에도 관련되어 있는 MYB2와 같은 모티브가 약 특이적인 발현을 주도했을 것으로 추정된다. 약 특이적인 프로모터는 알러겐이나 교배육종에 필요한 교배모본 형성 또는 GM작물에서의 화분비산 방지등의 다양한 목적으로 벼나 배추 과 등에서 지속적으로 연구하고 있다. 나리의 경우 봉오리 상태로 수확을 해서 판매가 진행되는데 이 과정에서 개약이 되면서 상품성이 떨어지는 경우가 많아 개약과정을 억제하거나 처음부 터 화분형성이 안되게 육종할 경우 상품성이 향상될 수 있다. 비록 형질전환이 어렵기는 하나 나리에서 분리한 gALCHS7 프로 모터를 이용하여 형질전환을 시도할 경우 보다 우수한 나리 품종을 만들어 활용할 수 있을 것으로 보이며 단자엽에서 분리한 프로모터가 쌍자엽에서도 잘 작동한 것으로 보아 배추나 기타 약 제어에 필요한 작물에 충분히 응용 가능할 것으로 생각된다.
안토시아닌은 화색을 결정하는 주요한 색소중의 하나이다. ALCHS2 유전자의 5' upstream 1584 bp 염기서열을 아카풀코 나리(Lilium Oriental hybrid cv. Acapulco)로부터 분리하였다. 컴퓨터 데이터 베이스 (GeneScan, AtPAN))는 CAATBOX1박 스와 조직특이적인 요소를 포함한 전사인자 결합부위를 예측하 였다. gALCHS7 프로모터와 GUS 연결체를 페튜니아(드림레드) 에 삽입했을 때 10개 형질전환체 모두가 약에서 GUS 발현을 나타냈으며 이중 5개 계통에서는 자방에서 약하게 GUS발현을 보였다. 동일한 단계에서 잎과 꽃잎에서는 어떤 구별되는 시그널 이 나타나지 않았다. 프로모터의 작동부위를 구명하기 위해서 형질전환체 6번의 약과 자방을 dark-field분석을 위해 파라핀에 고정하였다. 봉오리가 1cm일때는 약에서는 GUS시그널이 관찰 되지 않았지만 자방에서는 희미한 발현을 보였다. 개약직전에 pollen, endothecium, epidermis에서 GUS가 강하게 발현되었 다. 4개의 형질전환체의 각각 기관의 fluoromatric GUS분석은 34번을 제외한 모든 라인의 약이 35S CaMV프로모터에 비해 높은 GUS활성을 보였다. cis-acting elements에 따른 절단시리 즈를 통해 최소부위(gALCHS7-7, 270 bp)가 비록 원래 프로모터 에 비해 약하긴 하지만 약에서만 GUS발현을 나타내는 것을 확인하였다.
본 연구는 농촌진흥청 어젠다 과제(과제번호 PJ010881)연구 비에 의해 이루어진 것임
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The characterization of anther-specific promoter encoding chalcone synthase isolated from Lilium Oriental hybrid cv. Acapulco
Korean. J. Breed. Sci.. 2017;49(3):190-199.   Published online September 1, 2017
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Korean. J. Breed. Sci.. 2017;49(3):190-199.   Published online September 1, 2017
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The characterization of anther-specific promoter encoding chalcone synthase isolated from Lilium Oriental hybrid cv. Acapulco
Image Image Image Image Image Image
Fig. 1 Nucleotide sequence from lily genomic clone gALCHS7. Only 1584 nt of the genomic sequence are shown. The start codon of ALCHS2 are shown in upper case (M). Computer deduced putative cis-elements are boxed (http://atpan.itps.ncku.edu.tw/index.php). Putative anther specific promoter region (gALCHS7-7) was amplified with forward primer (underlined) and the same reverse primer of gALCHS7 promoter.
Fig. 2 Histochemical localization of GUS activity transgenic petunia plants transformed with gALCHS7::uidA. A, non-transgenic plant; B, transgenic petunia harboring the gALCHS::uidA gene; C, transgenic petunia harboring the pBI121. An, anther ; Ov, ovule ; Pe, petal.
Fig. 3 Dark-field analysis of a non-transgenic (A-E) and gALCHS7::uidA transformed plants(#6) (F-J). GUS expression at bud stage of 1 cm in size (A and F), before anthesis (B and G), and mature pollen (C and H). Longitudinal sections of ovule at bud stage of 1 cm in size (D and I), and before anthesis (E and J). Scale Bars, 1cm.
Fig. 4 Changes in GUS activity drived by gALCHS7::uidA promoter and pBI 121 (35S CaMV::uidA) in petunia anther and ovules. GUS activity is averaged from three independent experiments. Vertical bars indicates standard deviation. The letters above colums indicate statistically significant differences at P<0.05 by SAS software (version SAS 8.2).
Fig. 5 Histochemical localization of GUS activity in petunia plants under gALCHS7-7 promoter. A, non-transgenic plant; B-C, transgenic petunia harboring the gALCHS7-7::uidA gene (B, line 16 and C, line 21). An, anther ; Ov, ovule ; Pe, petal
Fig. 6 Representative results of gALCHS7-7 promoter through dark-field analysis. Stained anthers from Fig. 5 were embedded in paraffin and sectioned. A-B; anther from non-transgenic plants, C-F; transgenic petunia harboring the gALCHS7-7::uidA gene (C-D, line 16 and E-F, line 21). Scale Bars, 1cm.
The characterization of anther-specific promoter encoding chalcone synthase isolated from Lilium Oriental hybrid cv. Acapulco