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Korean. J. Breed. Sci. : Korean Journal of Breeding Science

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β-카로틴 강화벼의 분자생물학적 특성과 안전성 평가

Molecular Biological Characteristics and Biosafety Assessment for β-carotene Biofortified Transgenic Rice

Korean Journal of Breeding Science 2015;47(1):29-38.
Published online: February 28, 2015

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*Corresponding author (E-mail: thryu@korea.kr, Tel: +82-63-238-4711, Fax: +82-63-238-4704)
• Received: January 30, 2015   • Revised: February 24, 2015   • Accepted: March 3, 2015

© The Korean Society of Breeding Science

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  • The β-carotene biofortified transgenic rice was developed by transforming rice cv. Nakdongbyeo with phytoene synthase (Psy) and carotene desaturase (Crt I) genes isolated from Capsicum and Pantoea. The aim of this study was to perform molecular characterization of rice transformants of T5-T7 generation harboring Psy and Ctr I genes driven by endosperm specific globulin promoter for biosafety evaluation of β-carotene biofortified transgenic rice. The structure and sequence of T-DNA in the transformation vector and the insertion sites, flanking sequences and generational stability of inserted T-DNA in transgenic rice lines were analyzed. The transformation vector consisted of right border, MAR gene, carotenogenic genes unit, herbicide resistance selectable marker unit, MAR gene and left border in sequential order. T-DNA was introduced at the position of 30,363,938-30,363,973 bp of chromosome No. 2 by adaptor-ligation PCR. Stable integration of T-DNA and stable expression of bar gene was confirmed in T5 to T7 generations. It was also confirmed that the backbone DNA of transformation vector containing antibacterial gene was not present in the genome of β-carotene biofortified transgenic rice. HPLC analysis confirmed that carotenoids were consistently detected through T5-T7 generations.
콩, 옥수수, 벼 등 세계 주요 식량 작물에 다른 종에서 분리 한 제초제저항성, 해충저항성 등의 유용유전자를 도입하여 유 전자변형(Genetically Modified, GM) 작물로 개발하여 환경 변화에 따른 식량문제를 해결할 수 있는 대안으로 대두되고 있다. GM 작물은 노동력과 생산비용을 줄이면서 수확량을 높일 수 있어 상업적으로 증가 추세이다(Brookes & Barfoot 2007). GM 작물 재배면적은 1996년 170만ha였으나, 2014 년에는 100배 이상 증가된 1억 7,530만 ha로 확대되었다. 또 한, 전세계적으로 GM 작물은 1,800만 명의 생산자가 재배하 고 있다(James 2013). 국내에도 사료 또는 식용을 목적으로 옥수수, 면실, 대두 등 많은 양의 GM 곡물들이 수입되고 있 다(KBCH 2014). 그러나, 일부 GM 작물의 환경 및 생태계 위해성과 식품 섭취에 따른 인체 영향 및 안전성에 대한 우려 가 제기되고 있다(De Vries & Wackernagel 2004).
벼는 아시아 국가를 포함해 세계적으로 중요한 식량작물이 다. 벼는 여러 단계의 도정 과정 중에 Fe, Zn 등의 미네랄과 비타민 A와 비타민 E 등의 여러 가지 필수 미량영양소가 손 실될 우려가 있다(Juliano & Bechtel 1985). 이러한 쌀의 영 양학적 문제를 개선하기 위한 방법으로 쌀의 배유에 프로비 타민 A인 β-carotene(β-카로틴)이 강화된 GM벼인 황금쌀 (Golden Rice)이 개발되었다. 황금쌀은 최초로 자포니카 벼 품종에 수선화(Narcissus pseudonarcissus) 유래의 phytoene synthase (Psy) 유전자와 세균 Erwinia uredovora 유래의 carotene desaturase (Crt I) 유전자들을 형질전환하여 개발되 었으며(Ye et al. 2000), 이어서 인디카 벼 품종에서도 형질전 환 벼가 개발되었다(Hoa et al. 2003). Paine et al. (2005)에 의해서 옥수수 유래 Psy 유전자가 도입된 2세대 황금쌀이 개 발되었으며, 1세대 황금쌀에 비해 카로티노이드(carotenoid) 함량이 23배 정도 증가되었다(Paine et al. 2005). 국내에서도 고추(Capsicum annuum) 유래의 Psy 유전자를 이용하여 carotenoid 대사 관련 다중유전자를 동시발현을 통한 β-카로 틴 강화벼가 개발되었다(Ha et al. 2010).
현재까지 국내에서는 상업적으로 재배가 승인된 GM 작물 은 없으나, 대학, 기업 및 국공립연구소에서 잔디, 벼, 고추, 감자 등의 많은 유용 GM 작물들이 개발되고 있으며, 제초제 저항성 GM 잔디, 바이러스 내병성 GM 고추, 레스베라트롤 생합성 GM 벼 등이 안전성 평가를 통해 상업화를 위한 단계 를 준비 중에 있다(Moon et al. 2013, Oh et al. 2014). GM 작물들의 실용화를 위해서는 환경에 미치는 잠재적 위험성, 즉 도입 유전자들이 표적 및 비표적 생물체로 전이될 가능성 과 잡초화, 생태계 교란 등에 대한 환경위해성 및 독성, 알레 르기와 영양 성분 평가 등의 관한 식품안전성뿐만 아니라, GM 작물들이 세대 진전에 따라 도입유전자의 발현정지, 유 전자 변이 등이 나타날 수 있어 GM 작물의 상업에 위해서는 세대 진전에 따른 도입 유전자의 후대안정성 평가가 필수적 이다(Oh et al. 2014, Xia et al. 2009).
본 연구에서는 β-카로틴 강화벼의 도입 유전자들에 대한 복수 세대 동안의 안정성 분석을 위해서 β-카로틴 강화벼에 도입된 목적 유전자들인 Crt I와 Psy, 개시인자, 종결인자, 선 발마커(bar) 및 MAR (Marix attachment region) 유전자들의 3세대에 대한 서던 분석과 도입 위치 염기서열 분석, 생육시 기별 PAT (Phosphinothricin acetyltransferase) 발현 및 최종 목적 산물인 카로티노이드 발현량 분석을 수행하였다. 본 시 험을 통해 국내 개발된 GM 벼의 분자생물학적 특성뿐만 아 니라, 도입된 목적 형질에 대한 안정성 평가를 하고자 하였다.
도입유전자의 인접서열 분석
β-카로틴 강화벼의 게놈에 도입된 T-DNA의 인접 염기서 열을 확인하고자 Ha et al. (2010)의 방법에 따라 수행하였다. β-카로틴 강화벼의 T5와 T7세대의 게놈 DNA를 Hae III로 절단하고, AD1와 AD2으로 구성된 adaptor를 접합하였다. 두 번째 단계로 adaptor에 대한 A1 (5’-TAATACGACTCACTATAG- 3’)과 A2 (5’-GACTCACTATA GCAATTAAC-3’) 의 특이 프라이머를 제작하고, T-DNA 오른쪽 경계(Right border) 에 대해서 RB1 (5’-GCTTGCTGAGTGGCTCCTTC-3’)과 RB2 (5’-CAAACGTAAAACGGCTTGTCC-3’)를 왼쪽 경 계(Left border)에 대해서는 LB1 (5’-GTCAGGAGCTCGAATTCAGT- 3’)와 LB2 (5’-CATTAAAAACGTCCGCAATG-3’) 의 프라이머를 제작하여 단계별로 PCR을 수행하였다. 이후 증 폭된 PCR산물을 1% 한천 겔에 전기영동하고 Gel Extraction Kit (QIAquick Gel Extraction Kit, Qiagen, Valencia, CA, USA)를 이용하여 정제 후, pGEM T-easy 벡터(pGEM T-easy vector systems, Promega, Madison, WI, USA)에 삽입하여 염기서열을 확인하였다. 확인된 염기서열은 NCBI BLAST (http://blast.ncbi.nlm.nih.gov/Blast.cgi) 분석을 이용하여 동 일성 분석을 수행하였다.
Genomic DNA 분리 및 서던 블롯 검정
β-카로틴 강화벼(β-PAC T5, T6, T7)와 모품종인 낙동벼 식물체의 잎을 각 1 g씩, 막자 사발에서 액체질소를 이용하여 마쇄한 후, DNeasy Plant kit (Qiagen, Valencia, CA, USA)로 genomic DNA를 분리 추출하였다. NanoDrop Spectrophotometer ND-1000 (NanoDrop Technologies Inc, Wilmington, DE, USA)을 이용하여 260/280 nm 값이 1.8-2.0 사이인 DNA를 실험에 이용하였다. β-카로틴 강화벼의 도입유전자 특성 검 정을 위해서 추출한 genomic DNA 10μg를 제한효소 Bam HI, Xho I 및 EcoR I 으로 절단하고 1% 한천 겔에서 전기 영동한 후, 변성 과정을 수행하였다. 변성된 한천 겔의 DNA를 nylon membrane (Hybond-N+, Amersham, Uppsala, Sweden) 에 전이시키고, membrane의 DNA 단편들을 UV-crosslink (1,200 × μJ/cm2)로 고정한 후 hybridization buffer (0.5 M Na2PO4, pH 7.2, 1% BSA, 7% SDS, 1 mM EDTA, 10 mg/mL salmon sperm testicle DNA)로 1시간 동안 전처리하였다. 혼 성화는 Random primer DNA labeling kit (Takara Bio Inc., Shiga, Japan)를 이용하였고, 방사선 프로브인 α-32P로 표지 하여, 65°C에서 16-18시간 동안 혼성화하였다. 도입 유전자 의 도입안정성 검정용 프로브는 특이적인 프라이머를 이용하 여 카로티노이드 생성 유전자(Crt I, Psy), 개시인자(35S, Glb), 종결인자(NOS, PinII), 선발마커(bar) 및 MAR 유전자별로 PCR로 증폭한 후 추출 및 정제하여 사용하였다(Table 1). Backbone DNA 유무 검정용 probe는 항생제인 spectinomycin 의 aadA (SPM) 유전자 부위를 PCR로 증폭한 후 추출 및 정 제하여 사용하였다(Table 2). Hybridization이 끝난 membrane 은 세척 용액(1st solution, 2X SSC, 0.1% SDS; 2nd solution, 1X SSC, 0.1% SDS; 3rd solution, 0.2X SSC, 0.1% SDS) 으로 세척한 후 Bio-imaging analyzer (BAS-2000, Fuji Photo film, Tokyo, Japan)로 분석하였다.
Table 1.
Primers list used for Southern blot analysis.
Table 1.
Probe Primer Primer sequence Product Size (bp)

Psy Forward 5'-ATTACTCCGGCGGCCTTAGA-3' 419
Reverse 5'-TCGGATAGACCTGCCTGTGC-3'
Crt I Forward 5'-CTTCGCGCCGCACCTCAACTG-3' 543
Reverse 5'-GTAACGCGGGCCGAAACAAACC-3'
bar Forward 5'-CTGCACCATCGTCAACCACT-3' 505
Reverse 5'-GGAAATTCGGGGTCATCAGA-3'
Glb Forward 5'-GGGACCCGTTGTATCTGGTT-3' 600
Reverse 5'-TCGGATGGGGTTTTATAGGG-3'
Pin II Forward 5'-TGGGCATCAAAGTTGTGTGT-3' 593
Reverse 5'-TTGACAATCCATTCGTTTCT-3'
NOS Forward 5'-ATTGAATCCTGTTGCCGGTC-3' 237
Reverse 5'-AGCTTGATATCGAATTCCCG-3'
35S Forward 5'-ACGCAGCAGGTCTCATCAAG-3' 554
Reverse 5'-ATCGCAATGATGGCATTTGT-3'
MAR Forward 5'-TGCCGATGAAAGGATATAACA-3' 450
Reverse 5'-GCATCAAGCTGGCCTAAACTTA-3'
Backbone DNA의 도입 유무 PCR 검정
β-카로틴 강화벼(β-PAC T7)의 genomic DNA에 유전자 운반체의 T-DNA 이외의 backbone DNA의 삽입유무 확인을 위한 유전자 운반체의 유전정보를 바탕으로 β-카로틴 강화벼 의 Backbone DNA (pSB11 vector) 확인용 프라이머를 제작 하였다(Table 2). PCR 검정을 위하여 dNTP (10 mM) 4 μL, 10X PCR buffer 4 μL, 프라이머 각 20 μM, f-Taq DNA polymerase 1 unit (Solgent, Daejeon, Korea), 주형 DNA로 pSB11-β-PAC vector 10 ng과 β-카로틴 강화벼(β-PAC T7) DNA 1 μg를 추가한 후 최종 반응 부피를 40 μL로 하였다. PCR 반응은 PTC-100 Thermal cycler (MJ Research, Waltham, MA, USA)를 이용하여 1회 반복(95°C, 5분), 35회 반복(9 5°C, 40초 - 60°C, 40초 - 72°C, 60초), 1회 반복(72°C, 5분) 반응을 순차적으로 실시하였다. 산물들을 1% 한천 겔에서 전 기 영동한 후 UV조사로 확인하였으며, Gel Extraction kit (QIAquick Gel Extraction Kit, Qiagen, Valencia, CA, USA) 를 이용하여 정제하고 pGEM T-easy vector (pGEM T-easy vector systems, Promega, Madison, WI, USA)에 삽입하여 정확한 염기서열 정보를 확인하였다.
Table 2.
Primers list used for Backbone DNA detection PCR analysis.
Table 2.
Probe Primer Primer sequence Product Size (bp)

pSB11 131-1300 131-1300 Forward 5'-ATTACTCCGGCGGCCTTAGA-3' 1,170
131-1300 Reverse 5'-TCGGATAGACCTGCCTGTGC-3'
pSB11(SPM) 1304-2092 1304-2092 Forward 5'-CTTCGCGCCGCACCTCAACTG-3' 789
1304-2092 Reverse 5'-GTAACGCGGGCCGAAACAAACC-3'
pSB11 2101-3400 2101-3400 Forward 5'-CTGCACCATCGTCAACCACT-3' 1,300
2101-3400 Reverse 5'-GGAAATTCGGGGTCATCAGA-3'
pSB11 3501-5100 3501-5100 Forward 5'-GGGACCCGTTGTATCTGGTT-3' 1,600
3501-5100 Reverse 5'-TCGGATGGGGTTTTATAGGG-3'
pSB11 5101-6100 5101-6100 Forward 5'-TGGGCATCAAAGTTGTGTGT-3' 1,000
5101-6100 Reverse 5'-TTGACAATCCATTCGTTTCT-3'
RAc1 RAc1 Forward 5'-ATCACTGCCTTGCTCCTAGC-3' 350
RAc1 Reverse 5'-GTACTCAGCCTTGGCAATCC-3'
도입유전자의 단백질 발현 확인
β-카로틴 강화벼에 PsyCrt I 유전자와 함께 도입된 Phosphinothricin acetyltransferase (PAT, bar) 유전자의 단 백질 발현량을 확인하기 위하여 웨스턴 블롯(Western blot) 검정을 실시하였다. β-카로틴 강화벼(β-PAC T5, T6, T7)와 낙동벼의 잎을 각 0.5 g씩을 취하고, 막자사발에서 액체질소 와 함께 분말화하였다. 분말화된 시료에 단백질 추출용액 (0.5 M tris-HCl, 10 mM EDTA, 0.7 M sucrose, 0.1 M KCl, protease inhibitor cocktail)을 첨가한 후, 원심분리 (4°C, 15,000 rpm, 20분)하여 단백질을 분리, 추출한 후 bovine serum albumin (BSA)을 기준으로 protein assay reagent (Bio-Rad, Hercules, CA, USA)를 이용하여 단백질을 정량하였다. 추출한 단백질 을 30 μg씩 10% SDS-PAGE 겔에서 전기영동 후, nitrocellulose membrane (Bio-Rad, Hercules, CA, USA)에 전이하였다. 전이된 membrane을 blocking 용액(5% non-fat skim milk, 0.02% sodium azide, TBST용액)으로 실온에서 2시간 동안 블로킹한 후, PAT antibody(1차 항체, 1:1000)로 4°C에서 12 시간 동안 처리하였고, anti-rabbit IgG-HRP antibody(2차 항 체, 1:2000)로 4°C에서 10시간 동안 처리하였다. 각 항체를 처리한 후, TBST 용액으로 10분간 4번씩 세척하였다. 면역 반응된 단백질은 ECL western blotting detection reagent (GE Healthcare, Piscataway, NJ, USA)를 이용하여 검출하였다.
PAT 단백질의 농도를 정량하기 위하여 β-카로틴 강화벼 (β-PAC T5, T6, T7)와 낙동벼의 성장 시기별로 잎을 채취하 여 각 시료들을 마쇄한 후 PBST용액과 함께 균질화 한 후 저 온처리(얼음에서 5분) 및 원심분리(4°C, 15,000 rpm, 5분)하여 단백질을 분리, 추출한 후 PAT/bar ELISA kits (EnviroLogix LibertyLink, Portland, OR, USA)를 이용하여 ELISA분석을 실시하였다. 모든 시료들은 상온에서 2시간 반응한 후 ELISA reader (Multiskan EX, Thermo Scientific, Waltham, MA, USA)를 이용하여 450 nm 파장에서 흡광도를 측정하였다 (Oh et al. 2014).
카로티노이드 분석
β-카로틴 강화벼에 도입된 PsyCrt I 유전자의 발현을 통한 최종 산물인 카로티노이드 함량을 확인을 위하여 고속액 체 크로마토그래피(High performance liquid chromatography, HPLC)를 실시하였다(Kim et al. 2010). β-카로틴 강화벼(β -PAC T5, T6, T7)와 낙동벼의 현미 시료 0.3 g에 0.1% ascorbic acid을 함유한 에탄올(w/v), 3 mL을 넣고, 85°C 항 온수조에 5분 처리한 후, 카로티노이드를 추출하였다. 여기에 80% potassium hydroxide, 120 μL를 넣고, 85°C 항온수조 에 10분간 처리하여 비누화한 후, 즉시 실온으로 냉각하였다. 냉각 후, 탈이온수 1.5 μL와 내부 표준물질로 25 μg/mL 농 도의 β-Apo-8'-carotenal 50 μL과 hexane 1.5 mL을 혼합한 후, 원심분리(3600 rpm, 5분)하여 카로티노이드를 추출하였 다. Hexane을 이용한 카로티노이드 추출은 2번 수행하였다. 추출액은 질소가스를 이용하여 농축시킨 후, dichloromethane/ metanol (50:50, v/v)에 다시 녹여 HPLC로 분석하였다. 카로 티노이드는 photodiode array 검출기가 장착된 Agilent 1100 HPLC (Agilent Technologies, Massy, France)를 사용하여 450 nm 검출파장에서 측정하였다. 컬럼은 C30 YMC column (250 × 4.6 mm, 3 μm; Waters Corporation, Milford, MA, USA) 을 사용하였고, 이동상 A는 10 mM ammonium acetate를 함 유한 92% methanol, 이동상 B는 methyl tert-butyl ether를 사용하였다. 용매구배는 A: 83%, B: 17%로 시작하여, 23분 에 A: 70%, B: 30%, 29분에 A: 59%, B: 41%, 35분에 A: 30%, B: 70%, 40분에 A: 30%, B: 70%, 44분에 A: 83%, B: 17%, 55분에 A: 83%, B: 17%로 하여 분석하였다. 용매 유속은 1 μL/min 하였으며, 컬럼 온도는 40°C로 유지하였다. 카로티노이드 표준용액으로 Lutein, α-carotene, β-carotene, zeaxanthin (CaroteNature; Lupsingen, Switzerland)을 이용 하여 얻은 검량선으로부터 카로티노이드를 정량하였고, 내부 표준물질로는 β-Apo-8'-carotenal (Sigma Chemical Co. St. Louis, MO, USA)을 사용하였다.
도입 유전자의 도입위치 및 주변 염기서열 분석
본 시험에 공시한 β-카로틴 강화벼는 카로티노이드 대사관 련 유전자들인 고추(Capsicum annuum cv. Nockwang) 유래 Psy 와 세균(Pantoea ananatis)에서 분리된 Crt I 유전자를 2A 시스템(동시발현기술)을 이용하여 벼에 도입하여 개발되 었다(Ha et al. 2010).
β-카로틴 강화벼 종자에서 β-카로틴의 함량은 모품종인 낙동벼에 비해 9배 증가되었다. 벼의 종자에 카로티노이드 성 분이 증진된 β-카로틴 강화벼 복수 세대(β-PAC T5-T7)에 대한 도입 유전자들의 유전적 안정성 분석을 실시하였다. 우 선, 도입유전자의 분자생물학적 특성 분석은 β-카로틴 강화 벼(β-PAC T5, T7)의 T-DNA 삽입부위의 인접서열을 Ha et al. (2010)의 방법에 따라 단계적으로 수행하였다. 먼저, 추출 한 genomic DNA를 제한효소 Hae III로 처리한 후, PCR을 수행하고 증폭된 Right border (RB)와 Left border (LB)의 PCR 산물은 pGEM T-easy 벡터에 클로닝하여 염기서열을 분 석하고, 이들 염기서열에 대하여 NCBI BLAST 검색을 실시 하여 확인한 인접서열을 Fig. 1C, 1D에 나타내었다. BLAST 검색 결과 운반체 내의 T-DNA 이외의 염기는 확인되지 않아 서, T-DNA만이 벼의 게놈에 삽입되었다. 또한, T-DNA의 인 접서열에 대한 게놈의 유사성 분석과 삽입위치를 확인한 결 과, 유사성 있는 염기서열은 확인되지 않았으며, T-DNA의 삽 입부위는 도입유전자의 RB 및 LB의 인접염기서열을 Gene Bank (NCBI)에서 Oryza sativa (japonica cultivar-group) 게 놈 데이터베이스에 근거하여 BLAST한 결과, RB 쪽의 염기 서열은 벼 염색체 2번 중에서 30,363,695-30,363,938번, LB 쪽은 염색체 2번 중 30,363,973-30,364,275번 사이의 염기서 열과 일치하였다. BLAST 염기서열 분석 결과를 바탕으로 도 입유전자의 삽입 위치를 확인한 결과, 벼 염색체 2번 중 30,363,938번과 30,363,973번 사이의 단일 부위에 도입유전 자가 삽입되어 있는 것을 확인하였다. 따라서 β-카로틴 강화 벼의 T4와 T6의 주변 염기 서열 분석이 서로 일치하는 결과 를 확인하였으며, β-카로틴 강화벼의 도입유전자가 벼 2번 염 색체 내에서 intergenic한 상태로 안정적으로 유지되고 있음 을 확인하였다(Table 3).
Table 3.
Junction sequence analysis of T-DNA in β-carotene biofortified transgenic rice.
Table 3.
Pedigree Chromosome Junction sequence Junction gene & sequence

β-carotene biofortified rice 2 30,363,695-30,363,93830,363,973-30,364,275 Os02g072910030,361,505~30,362,937Os02g072930030,371,203~30,374,971
Fig. 1.
T-DNA junction sequences analysis. (A) Schematic representation of the pSB11-β-PAC vector used for rice transformation. pSB11-β-PAC vector contained a recombinant gene, Psy-2A-Tp-Crt I (β-PAC). The remaining construct components had the same configuration, consisting of the rice endosperm-specific globulin promoter (Glb), the 3'region of the potato proteinase inhibitor II gene (PinII) and the Bar expression cassette containing a 35S promoter/Bar coding region/3'region from the nopaline synthase gene (NOS). β-PAC entire expression cassette was flanked by a 5'-matrix attachment region (Mar) from the chicken lysozyme gene. (B) Secondary PCR products and junction sequences of T-DNA (C: Right border; D: Left border) for the β -carotene biofortified transgenic rice line flanking region. The genomic DNA was digested with Hea III, ligated to genome walking adapter to create different libraries, and used as template for PCR. The DNA molecular size markers are indicated on the left. RB: right border, LB: left border. Capital letters represents the flanking genomic sequences and lower case letters show the inner part of each border of T-DNA.
KJBS-47-29_F1.gif
도입 유전자 서던 분석
β-카로틴 강화벼의 도입 유전자(T-DNA)는 카로티로이드 생성 유전자(Crt I, Psy), 선별마커(bar), 개시인자(35S, Glb), 종결인자(NOS, PinII) 및 MAR 유전자로 구성되어있다(Fig 1A). 도입 유전자의 후대안정성 분석에 사용된 복수세대의 증진된 β-카로틴 강화벼(β-PAC T5-T7) 시료에서 도입 유전 자들인 카로티로이드 생성 유전자(Crt I, Psy), 선별마커(bar), 개시인자(35S, Glb), 종결인자(NOS, PinII) 및 MAR 유전자 에 대한 서던 분석을 수행하였다. β-카로틴 강화벼(β-PAC T5-T7)의 잎에서 genomic DNA를 분리 추출한 후, 각 10 ug 를 Bam HI, Xho I 및 EcoR I 제한효소로 처리하고 한천겔에 서 전기영동한 후, membrane에 전이하였으며, 양성대조군 (PC)으로는 pSB11-β-PAC을 500 pg을 사용하였다. β-카로 틴 강화벼 각 세대별(β-PAC T5-T7)로 개시인자인 Glb35S 유전자를 이용하여 서던 분석을 수행한 결과, Glb 개시 인자(Xho I 절단)는 음성대조군인 낙동벼와 T4에서 T6세대 까지의 β-카로틴 강화벼에서 검출되었으며, 전체적으로 끌리 는 양상을 보였다. 이는 Glb 개시인자가 벼 내재 글로블린 유 전자에서 유래되었기 때문인 것으로 사료된다. 35S 개시인자 (Bam HI 절단)는 β-카로틴 강화벼의 3세대 동안 안정적으로 동일한 위치인 12 kb 크기의 한 개 밴드가 검출되었으며, 낙 동벼에서는 검출되지 않음을 확인하였다(Fig. 2A, E). β-카로 틴 강화벼(β-PAC T5-T7)에 대하여 β-카로틴을 생산하는 Crt I과 Psy 유전자(Xho I 절단)를 프로브로 이용하여 서던 분석 결과, 각 세대별로 12 kb 크기의 밴드인 한 카피의 유전자가 도입되었고 안정적으로 유전자가 유지되고 있음을 확인하였 다(Fig. 2B, C). β-카로틴 강화벼의 β-PAC T5-T7 세대별로 도입유전자 종결인자인 PinIINOS 유전자의 서던 분석(각 각 Bam HI 절단) 결과, 12 kb 크기의 한 카피의 유전자가 도 입되었고, T5-T7 세대별로 안정적으로 유전자가 유지되고 있 음을 확인하였다(Fig. 2D, G). β-카로틴 강화벼의 각 세대별 (β-PAC T5-T7)로 선발마커인 bar 유전자(Bam HI 절단) 및 MAR 유전자(EcoR I 절단)를 프로브로 이용하여 서던 분석 을 수행한 결과, bar 유전자는 β-카로틴 강화벼에서만 12kb 크기의 한 카피 유전자가 도입되었음을 확인하였고 낙동벼에 서는 검출되지 않았다. MAR 유전자 서던 분석 결과, 2개의 밴드가 검출되었는데, 이는 형질 전환된 T-DNA의 양끝에 두 개의 MAR 유전자가 도입됨을 확인하였다(Fig. 2F, H). 모든 도입 유전자들의 서던 분석 결과, 카로티노이드 생성 인자 (Crt I, Psy), 개시인자(35S, Glb), 종결인자(NOS, PinII), 선 별마커(bar) 및 MAR 유전자들이 본 실험에 사용된 β-카로틴 강화벼에 단일 카피로 도입되었고, 도입 유전자가 β-카로틴 강화벼 T5에서 T7세대까지 안정적으로 발현됨을 확인할 수 있었다.
Fig. 2.
Genomic Southern blot analyses of β-carotene biofortified transgenic rice. Genomic DNAs from leaf tissues were digested with Xho I, EcoR I and BamH I and hybridized with probe (A) Glb, (B) Crt I, (C) Psy, (D) PinII, (E) 35S, (F) Bar, (G) NOS, and (H) MAR. The DNA molecular size markers are indicated on the left. PC, PC means pSB11 vector carrying β-PAC recombinant genes as positive controls, respectively; Nakdong, non-transgenic wild type rice as negative controls.
KJBS-47-29_F2.gif
Backbone DNA의 도입 유무 PCR 검정
β-카로틴 강화벼 형질전환에 사용된 유전자 운반체의 backbone DNA인 pSB11 벡터에서 Fig. 3A와 같이 서던 분석용 프로 브 및 PCR 검정용으로 디자인 및 프라이머를 제작하였고 (Table 2), β-카로틴 강화벼 T7의 게놈 DNA를 대상으로 Table 2의 프라이머 셋을 이용하여 PCR을 수행하였고, 그 결과, β-카 로틴 강화벼 형질전환에 사용된 pSB11-β-PAC (Backbone DNA 와 T-DNA)에서는 pSB11 131-1300, pSB11 1304-2092 (spectinomycin; spm), pSB11 2101-3400, pSB11 3500-5100, pSB11 5101-6100 등 각각의 부위에 따라 양성대조군(PC)인 Backbone DNA에서만 밴드가 검출되었고, β-카로틴 강화벼 T7에서는 벼 내재 유전자인 actin 유전자 밴드만 생성되어, 항생제(spectinomycin) 내성 유전자(aadA)를 포함한 Backbone DNA가 벼 게놈에 도입되지 않음을 검정하였다(Fig. 3B). β- 카로틴 강화벼 T5-T7의 각 계통에 대한 항생제 내성 마커 유 전자인 aadA를 서던 분석 결과, 양성대조군(PC)인 pSB11-β -PAC에서만 밴드가 검출되었고, β-카로틴 강화벼의 세대별 (β-PAC T5~T7)에서는 aadA가 삽입되지 않음을 확인하였 다(Fig. 3C).
Fig. 3.
Backbone DNA PCR and Southern blot analyses of β-carotene biofortified transgenic rice. (A) Schematic representation of the pSB11 vector used for PCR and Southern blot analyses. (B) Verification of Backbone DNA (pSB11) insert using PCR amplification of genomic DNA samples extracted from transgenic leaf tissues. (C) Genomic DNAs from leaf tissues were digested with Bam HI and hybridized with probe aadA (SPM). The DNA molecular size markers are indicated on the left. PC, PC means pSB11 vector carrying β-PAC recombinant genes as positive controls, respectively; Nakdong, non-transgenic wild type rice as negative controls.
KJBS-47-29_F3.gif
도입유전자 복수세대의 단백질 발현 확인
β-카로틴 강화벼에서 선발마커로 사용된 제초제 저항성인 PAT(bar 유전자) 단백질 발현을 검정하기 위하여 웨스턴 블 롯을 실시하였다. β-카로틴 강화벼(β-PAC T5-T7)와 낙동벼 의 잎에 대하여, 추출한 단백질을 10% SDS-PAGE 겔에서 전기영동 후, 전이된 nitrocellulose membrane에 PAT 항체를 이용하여 단백질 발현을 분석한 결과, β-카로틴 강화벼(β -PAC T5-T7)에서만 21 kDa에 특이적으로 단백질 밴드가 검 출되었으며, 대조구인 낙동벼에서는 단백질이 검출되지 않았 다(Fig. 4). 또한, β-카로틴 강화벼에서의 생육 시기(유묘기, 성장기, 출수기, 성형기)별로 PAT 단백질 발현량을 정량하기 위해서 ELISA (Enzyme-linked immunosorbent assay)법을 이용하여 분석한 결과, β-카로틴 강화벼 에서만 발현되었다. β-카로틴 강화벼 생육 시기(유묘기, 성장기, 출수기, 성형기) 별로 T5 세대는 각각 0.50 ± 0.01 mg/g, 0.19 ± 0.00 mg/g, 0.09 ± 0.00 mg/g, 0.06 ± 0.00 mg/g 였으며, T6 세대에서는 각각 0.41 ± 0.02 mg/g, 0.22 ± 0.00 mg/g, 0.11 ± 0.00 mg/g, 0.06 ± 0.00 mg/g 였으며, T7 세대에서는 각각 0.51 ± 0.01 mg/g, 0.21 ± 0.00 mg/g, 0.10 ± 0.00 mg/g, 0.06 ± 0.00 mg/g 으로 측정되었으며, 모품종인 낙동벼에서는 검출되지 않음을 확인하였다(Table 4).
Table 4.
PAT protein levels in Nagdongbyeo and β-carotene biofortified transgenic rice (β-PAC).
Table 4.
Sample PAT Concentration (mg/g)
Seedling stage Tillering stage Heading stage Ripening stage

Nagdongbyeo 0 (N.D.) 0 (N.D.)z 0 (N.D.) 0 (N.D.)
β-PAC T5 0.50 ± 0.01y 0.19 ± 0.00 0.09 ± 0.00 0.06 ± 0.00
β-PAC T6 0.41 ± 0.02 0.22 ± 0.00 0.11 ± 0.00 0.06 ± 0.00
β-PAC T7 0.51 ± 0.01 0.21 ± 0.00 0.10 ± 0.00 0.06 ± 0.00

zN.D. : Not detected

yValues are the average ± SD of triplicate measures

Fig. 4.
Expression analysis of PAT at the β-carotene biofortified transgenic rice and non-GM rice using western blot. Crude protein extracts were isolated from leaf tissue, separated on 10% SDS-PAGE, and then visualized through the immunoblotting process. M: Protein ladder, N: non-GM rice (Nakdong), β-PAC: β-carotene biofortified transgenic rice.
KJBS-47-29_F4.gif
카로티노이드 분석
β-카로틴 강화벼의 PsyCrt I의 단백질 발현 산물인 카 로티노이드를 β-카로틴 강화벼의 각 세대별로 수확한 종자에 서 HPLC법을 이용하여 분석한 결과, β-카로틴 함량은 β -PAC T5는 1.43 μg/g, β-PAC T6는 1.65 μg/g, β-PAC T7 는 1.43 μg/g 으로 세대 평균 1.50 μg/g으로 측정되었고, 이 는 모품종인 낙동벼의 0.16 μg/g에 비해 9.4배 함량이 증가되 었다. α-carotene와 zeaxanthin은 대조군인 낙동벼에서는 검 출되지 않았으나, β-카로틴 강화벼 세대에서는 각각 평균 0.39 μg/g와 0.24 μg/g이 검출되었다(Table 5). β-카로틴 강 화벼의 세대별(β-PAC T5-T7)로 안정적으로 도입 유전자의 단백질이 발현됨을 확인할 수 있었다.
Table 5.
Seed carotenoid content and composition of β-carotene biofortified transgenic rice.
Table 5.
Carotenoid (µg/g) Plants
Nagdongbyeo β-PAC T5 β-PAC T6 β-PAC T7

lutein 0.23 ± 0.08 0.32 ± 0.05 0.51 ± 0.18 0.72 ± 0.06
zeaxanthin N.D. 0.19 ± 0.02 0.23 ± 0.03 0.30 ± 0.01
α-carotene N.D. 0.37 ± 0.06 0.42 ± 0.11 0.37 ± 0.05
β-carotene 0.16 ± 0.03 1.43 ± 0.12 1.65 ± 0.64 1.43 ± 0.16

Data are expressed as mean (ug/g dry weight) ± SD from three independent experiments using individual T5~T7 mature seeds. N.D. : Not detected

GM작물의 상업화를 위해서는 도입 유전자의 안정성 평가 와 식품 및 환경에 대한 영향을 평가하기 위한 GMO 안전성 평가를 실시해야 하며, 분자생물학적 특성 평가와 도입유전자 의 작물 세포내 삽입으로 목적 형질에 대한 후대안정성 평가 는 필수항목이다(Oh et al 2014). 현재까지 국내에서 재배 승 인되어 상업화 된 GM벼 사례는 없지만, 안전성평가를 위해 GM벼의 분자생물학적 특성과 도입 유전자의 후대안정성을 검정한 여러 연구결과가 국내에서도 보고되었다(Ahn et al. 2014, Lee et al. 2014, Oh et al. 2014). 본 연구에서도 β-카 로틴 강화벼의 도입 유전자에 대한 벼 genomic DNA내의 도 입 위치와 안정성을 평가한 결과, 단일 부위에 intergenic한 상태로 β-카로틴 강화벼 도입유전자가 삽입되어 3세대 이상 의 복수세대에서 한 카피로 안정적으로 유지되고 있음을 확 인하였고, 선발 마커로 도입된 PAT 단백질의 발현 분석을 통 해 세대별 안정적으로 발현되고 있음을 확인하였다. β-카로 틴 강화벼의 목적 형질인 β-carotene은 모품종에 비해 함량이 9.4배 증가됨을 확인하였다.
β-카로틴 강화벼 도입유전자의 도입 위치와 안정성을 도입 위치 주변염기서열 분석과 서던 분석한 결과, 벼 염색체 2번 중 30,363,938번과 30,363,973번 사이의 단일 부위에 도입유 전가 도입되었으며, T5-T7 세대 동안 도입된 모든 유전자들 이 안정적으로 유지되고 있으며, 도입유전자의 운반체인 Backbone DNA (pSB11)는 β-카로틴 강화벼에 삽입되지 않 았음을 확인하였다. 선발 마커로 도입된 PAT 단백질의 발현 분석 결과에서도 T5-T7 세대별, 생육시기별로 안정적으로 발 현됨을 검정하였으며, 목적 형질인 β-카로틴 분석 결과에서 도 모품종인 낙동벼에 비해 9.4배 함량이 증가됨을 확인하였 다. 이상의 분석기법을 통해 복수세대에서 β-카로틴 강화벼 의 도입 유전자들이 안정적으로 유지되고 목적 단백질들이 안정적으로 발현되고 있음을 확인하였다.
본 연구는 농촌진흥청 국립농업과학원의 농업과학기술 연 구개발사업 지원(과제번호: PJ000960901)의 지원에 의해 수 행되었음.
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Molecular Biological Characteristics and Biosafety Assessment for β-carotene Biofortified Transgenic Rice
Korean. J. Breed. Sci.. 2015;47(1):29-38.   Published online March 31, 2015
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Molecular Biological Characteristics and Biosafety Assessment for β-carotene Biofortified Transgenic Rice
Image Image Image Image
Fig. 1. T-DNA junction sequences analysis. (A) Schematic representation of the pSB11-β-PAC vector used for rice transformation. pSB11-β-PAC vector contained a recombinant gene, Psy-2A-Tp-Crt I (β-PAC). The remaining construct components had the same configuration, consisting of the rice endosperm-specific globulin promoter (Glb), the 3'region of the potato proteinase inhibitor II gene (PinII) and the Bar expression cassette containing a 35S promoter/Bar coding region/3'region from the nopaline synthase gene (NOS). β-PAC entire expression cassette was flanked by a 5'-matrix attachment region (Mar) from the chicken lysozyme gene. (B) Secondary PCR products and junction sequences of T-DNA (C: Right border; D: Left border) for the β -carotene biofortified transgenic rice line flanking region. The genomic DNA was digested with Hea III, ligated to genome walking adapter to create different libraries, and used as template for PCR. The DNA molecular size markers are indicated on the left. RB: right border, LB: left border. Capital letters represents the flanking genomic sequences and lower case letters show the inner part of each border of T-DNA.
Fig. 2. Genomic Southern blot analyses of β-carotene biofortified transgenic rice. Genomic DNAs from leaf tissues were digested with Xho I, EcoR I and BamH I and hybridized with probe (A) Glb, (B) Crt I, (C) Psy, (D) PinII, (E) 35S, (F) Bar, (G) NOS, and (H) MAR. The DNA molecular size markers are indicated on the left. PC, PC means pSB11 vector carrying β-PAC recombinant genes as positive controls, respectively; Nakdong, non-transgenic wild type rice as negative controls.
Fig. 3. Backbone DNA PCR and Southern blot analyses of β-carotene biofortified transgenic rice. (A) Schematic representation of the pSB11 vector used for PCR and Southern blot analyses. (B) Verification of Backbone DNA (pSB11) insert using PCR amplification of genomic DNA samples extracted from transgenic leaf tissues. (C) Genomic DNAs from leaf tissues were digested with Bam HI and hybridized with probe aadA (SPM). The DNA molecular size markers are indicated on the left. PC, PC means pSB11 vector carrying β-PAC recombinant genes as positive controls, respectively; Nakdong, non-transgenic wild type rice as negative controls.
Fig. 4. Expression analysis of PAT at the β-carotene biofortified transgenic rice and non-GM rice using western blot. Crude protein extracts were isolated from leaf tissue, separated on 10% SDS-PAGE, and then visualized through the immunoblotting process. M: Protein ladder, N: non-GM rice (Nakdong), β-PAC: β-carotene biofortified transgenic rice.
Molecular Biological Characteristics and Biosafety Assessment for β-carotene Biofortified Transgenic Rice

Primers list used for Southern blot analysis.

Probe Primer Primer sequence Product Size (bp)

Psy Forward 5'-ATTACTCCGGCGGCCTTAGA-3' 419
Reverse 5'-TCGGATAGACCTGCCTGTGC-3'
Crt I Forward 5'-CTTCGCGCCGCACCTCAACTG-3' 543
Reverse 5'-GTAACGCGGGCCGAAACAAACC-3'
bar Forward 5'-CTGCACCATCGTCAACCACT-3' 505
Reverse 5'-GGAAATTCGGGGTCATCAGA-3'
Glb Forward 5'-GGGACCCGTTGTATCTGGTT-3' 600
Reverse 5'-TCGGATGGGGTTTTATAGGG-3'
Pin II Forward 5'-TGGGCATCAAAGTTGTGTGT-3' 593
Reverse 5'-TTGACAATCCATTCGTTTCT-3'
NOS Forward 5'-ATTGAATCCTGTTGCCGGTC-3' 237
Reverse 5'-AGCTTGATATCGAATTCCCG-3'
35S Forward 5'-ACGCAGCAGGTCTCATCAAG-3' 554
Reverse 5'-ATCGCAATGATGGCATTTGT-3'
MAR Forward 5'-TGCCGATGAAAGGATATAACA-3' 450
Reverse 5'-GCATCAAGCTGGCCTAAACTTA-3'

Primers list used for Backbone DNA detection PCR analysis.

Probe Primer Primer sequence Product Size (bp)

pSB11 131-1300 131-1300 Forward 5'-ATTACTCCGGCGGCCTTAGA-3' 1,170
131-1300 Reverse 5'-TCGGATAGACCTGCCTGTGC-3'
pSB11(SPM) 1304-2092 1304-2092 Forward 5'-CTTCGCGCCGCACCTCAACTG-3' 789
1304-2092 Reverse 5'-GTAACGCGGGCCGAAACAAACC-3'
pSB11 2101-3400 2101-3400 Forward 5'-CTGCACCATCGTCAACCACT-3' 1,300
2101-3400 Reverse 5'-GGAAATTCGGGGTCATCAGA-3'
pSB11 3501-5100 3501-5100 Forward 5'-GGGACCCGTTGTATCTGGTT-3' 1,600
3501-5100 Reverse 5'-TCGGATGGGGTTTTATAGGG-3'
pSB11 5101-6100 5101-6100 Forward 5'-TGGGCATCAAAGTTGTGTGT-3' 1,000
5101-6100 Reverse 5'-TTGACAATCCATTCGTTTCT-3'
RAc1 RAc1 Forward 5'-ATCACTGCCTTGCTCCTAGC-3' 350
RAc1 Reverse 5'-GTACTCAGCCTTGGCAATCC-3'

Junction sequence analysis of T-DNA in β-carotene biofortified transgenic rice.

Pedigree Chromosome Junction sequence Junction gene & sequence

β-carotene biofortified rice 2 30,363,695-30,363,93830,363,973-30,364,275 Os02g072910030,361,505~30,362,937Os02g072930030,371,203~30,374,971

PAT protein levels in Nagdongbyeo and β-carotene biofortified transgenic rice (β-PAC).

Sample PAT Concentration (mg/g)
Seedling stage Tillering stage Heading stage Ripening stage

Nagdongbyeo 0 (N.D.) 0 (N.D.)z 0 (N.D.) 0 (N.D.)
β-PAC T5 0.50 ± 0.01y 0.19 ± 0.00 0.09 ± 0.00 0.06 ± 0.00
β-PAC T6 0.41 ± 0.02 0.22 ± 0.00 0.11 ± 0.00 0.06 ± 0.00
β-PAC T7 0.51 ± 0.01 0.21 ± 0.00 0.10 ± 0.00 0.06 ± 0.00

N.D. : Not detected

Values are the average ± SD of triplicate measures

Seed carotenoid content and composition of β-carotene biofortified transgenic rice.

Carotenoid (µg/g) Plants
Nagdongbyeo β-PAC T5 β-PAC T6 β-PAC T7

lutein 0.23 ± 0.08 0.32 ± 0.05 0.51 ± 0.18 0.72 ± 0.06
zeaxanthin N.D. 0.19 ± 0.02 0.23 ± 0.03 0.30 ± 0.01
α-carotene N.D. 0.37 ± 0.06 0.42 ± 0.11 0.37 ± 0.05
β-carotene 0.16 ± 0.03 1.43 ± 0.12 1.65 ± 0.64 1.43 ± 0.16

Data are expressed as mean (ug/g dry weight) ± SD from three independent experiments using individual T5~T7 mature seeds. N.D. : Not detected

Table 1. Primers list used for Southern blot analysis.
Table 2. Primers list used for Backbone DNA detection PCR analysis.
Table 3. Junction sequence analysis of T-DNA in β-carotene biofortified transgenic rice.
Table 4. PAT protein levels in Nagdongbyeo and β-carotene biofortified transgenic rice (β-PAC).

N.D. : Not detected

Values are the average ± SD of triplicate measures

Table 5. Seed carotenoid content and composition of β-carotene biofortified transgenic rice.

Data are expressed as mean (ug/g dry weight) ± SD from three independent experiments using individual T5~T7 mature seeds. N.D. : Not detected