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

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해충저항성 유전자변형 벼(Agb0101) 유전자 이동성 평가

Assessment of gene flow from insect-resistant genetically modified rice (Agb0101) to non-GM rice

Korean Journal of Breeding Science 2017;49(3):180-189.
Published online: August 31, 2017

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*Corresponding Author (abychan@korea.kr, +82-63-238-4711, +82-63-238-4704)
• Received: June 16, 2017   • Accepted: July 31, 2017

© Korean Society of Breeding Science. All rights reserved.

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  • Genetically modified (GM) crops have been developed worldwide through the recombinant DNA technology and commercialized by global agricultural companies. Until now, GM crops have not been cultivated commercially in Korea. Commercialization of GM crops requires a compulsory assessment of environmental risk associated with the release of GM crops. This study was conducted to evaluate the frequency of pollen mediated gene flow from Bt transgenic rice (Agb0101) to japonica non-GM rice (Nakdongbyeo), indica non-GM rice (IR36), and weedy rice (R55). A total of 729,917, 596,318 and 230,635 seeds were collected from Nakdongbyeo, IR36, and R55, respectively, which were planted around Agb0101. Selection of the hybrids was determined by repeated spraying of herbicide and Cry1Ac1 immunostrip assay. Finally, the hybrids were confirmed by PCR analysis using specific primer. The hybrids were found in all non-GM rice and out-crossing ranged from 0.0005% at IR36 to 0.0027% at Nakdongbyeo. All of hybrids were located within 1.2 m distance from the Agb0101 rice plot. The meteorological elements including rainfall and temperature during rice flowering time were found to be important factors to determine rice out-crossing rate. Consideration should be taken for many factors like the meteorological elements of field and physiological condition of crop to set up the safety management guideline to prevention of GM crops gene flow.
1996년에 제초제내성 유전자변형(Genetically Modified, GM) 콩이 상업적으로 재배된 이후, 생명공학기술을 이용한 GM 작물의 재배면적은 매년 증가하고 있는 상황이다. 상업적인 재배가 20년된2015년에는 전 세계적으로 1억 7,970만 헥타르의 면적에서 GM 작물이 재배되었으며, GM종자의 시장규모도 153 억 달러에 이르며 세계종자시장의 20%에 육박하고 있다(James 2015). GM작물의 긍정적인 측면인 농약 사용 절감과 이에 따른 재배의 용이함, 생산량 증가에 따른 농가 소득 증대 등 다양한 잇점으로 GM작물의 개발과 재배는 증가되어왔으며(Owen 2000), 2000년대에 들어서면 아시아와 중남미 지역에서 증가되 고 있다(Lee et al. 2012).
GM 작물의 재배면적이 늘어나면서 농업환경 안전성에 대한 관심도 증가되고 있다. 그 중에 GM작물에 도입된 유전자의 교잡에 의한 일반 재배 품종의 유전자 오염, 잡초화 유발 및 야생종 감소 등에 따른 농업 생태계의 교란 우려도 있다(Conner et al. 2003). 이러한 유전자 교잡에 대한 사회적 우려와 관심이 높은 이유는 GM작물을 개발시에 선발을 위해 항생제 또는 제초 제 저항성 유전자를 주로 이용하기 때문이다. 이는 만약 GM작물 을 농경지 재배할 때 선발 유전자가 다른 작물 등으로 전달되면 목적유전자에 해당하는 항생제 또는 제초제 저항성을 갖는 슈퍼 잡초 또는 생태계 교란이 일어날수 있는 가능성이 존재하기 때문이다. 영국에서 연구에 의하면 사과, 당근, 사탕무 등 31종의 재배 품종화된 작물 중 30% 가량이 하나 혹은 그 이상의 야생종 과의 교잡이 가능하며(Raybould & Gray 1993), 네덜란드에서 는 연구된 42종의 식물 중 25% 정도가 교잡 가능성 있는 것으로 보고된 바 있다(Ellstrand et al. 1999). 국내에 수입되는 농산물 의 주요 생산국들로는 중국, 캐나다, 미국, 아르헨티나 등은 GM 작물을 대단위 면적에서 재배하고 있으므로, 국내에 수입되는 농산물에 GM작물이 혼입될 가능성이 존재하여 주요 GM작물의 환경 위해성 평가 기술 및 프로토코 개발의 필요성이 요구되고 있다. 또한, 최근 국내에서도 농업생명공학에 대한 집중적인 투자 와 연구를 통해 다수의 GM작물이 개발되었고 이를 활용하기 위해서는 GM 작물의 환경위해성 평가가 반드시 이루어져야 한다.
벼(Oryza sativa)는 동남아시아뿐만 아니라 밀, 옥수수와 더블 어3대 곡류 중 하나이며, 세계 인구의 절반이 주식으로 활용하고 있다(OECD 1999). 대표적인 자가수정 작물인 벼는 외부에 노출 된 화분의 생존시간이 짧아서 개화와 동시에 수분이 이루어지며, 일반적인 기상 조건에서 오전 11시경에 개화가 최고조에 도달한 다. 일반적인 벼의 자연 교잡율은 1% 내외로 보고되었으나, 제초제저항성 GM벼로부터 non-GM 벼로의 유전자 이동에 의 한 교잡율은 0.05~0.53%였으며, 이는 일반벼의 교잡율에 비해 1/2에서 최대 1/10낮은 수준으로 보고되었다(Messeguer et al. 2001). 제초제저항성 GM벼(Nam29/TR18)로부터 근연 잡초성 벼로의 교잡율은 0.001~0.046%로 보고되었다(Chen et al. 2004). 화분에 의한 벼의 교잡 가능 거리가 최대 30m이고, 일반 적인 벼 교잡율이 1% 이내이므로, 벼의 유전자 이동성은 농업 환경에 미치는 영향이 크지 않을 것으로 보고되었지만(Gealy et al. 2003), 같은 O. sativa에 속해 있는 잡초성벼와 재배품종 벼는 다양한 요인에 의한 유전자 교잡의 가능성이 있으며(Song et al. 2004), Nortai품종은 화분 공여체와 수여체의 개화기 중복 여부 및 공여체의 개화기간 연장 및 등에 따라 최대 52%까지의 교잡율이 확인된 바 있다(Langevin et al. 1990). 국내에서 2009 년도에 수행된 비타민 A강화벼의 유전자 이동성 평가 결과, 화분공여의 30cm에 위치한 낙동벼에서 9.12%의 높은 유전자 이동성이 보였고(Bae et al. 2013), 2010년부터 2014년까지 병저항성GM벼, 가뭄저항성 GM벼, 및 레스베라트롤 생합성벼 등의 실험 재배도 이루어졌지만 아직까지 국내에서는 GM작물 의 상업재배는 이루어지지 않고 있다(Oh et al. 2014).
본 연구에서는 토양 세균인 B. thuringiensis에서 유래한 해충 저항성 유전자(mCry1Ac1)를 낙동벼에 도입시킴으로써 혹명나 방(Cnaphalocrocis medinalis)과 두줄꼬마 밤나방 (Naranga aenescens), 벼줄점팔랑나비(Parnara guttata)에 대하여 살충성 을 나타내는 해충저항성 Bt벼(Shin et al. 2009)의 환경위해성 평가 항목 중 유전자 이동성에 대한 평가를 수행하였다. 본 연구 를 통해 국내 개발 GM벼의 환경위해성 평가 자료를 생산뿐만 아니라, GM작물의 환경위해성 평가 및 안전관리 지침서 작성에 활용하고자 하였다.
공시 재료
해충저항성 Bt 벼(Agb0101)는 토양 세균인 B. thuringiensis에 서 유래한 mCry1Ac1 유전자가 rbcS 프로모터와 pinII 터미네이터 에 의해 발현되는 운반체(Fig. 1)로 낙동벼에 형질전환되었고, 선발마커로 글루포시네이트(바스타, glufosinate ammonium) 제초제저항성 유전자(bar)가 함께 도입되어 있어 유전자 이동에 따른 교잡개체 선발에 이용하고자 하였다. 해충저항성 Bt 벼 (Agb0101)의 이벤트 계통(Cr7-1-9-1-1-4-1)과 선행 연구를 통 해 개화기가 유사한 잡초성벼(R55)와 자포니카 품종(낙동벼), 인디카 품종(IR36)를 각각 파종하고 포장에 이앙하기 전까지 온실에서 재배하였다.
Fig. 1
Schematic representation of the inserted vector used for rice transformation. The arrows indicate primer (Table 1) sites and the direction of synthesis. RB: right border; MAR: matrix attachment region; rbsS: rice rbcS promoter; TP: rice rbcS transit peptide; mCry1Ac1: B. thuringiensis mCry1Ac1 gene; PinII: potato proteinase inhibitor II gene; 35S: CaMV 35S promoter; PAT: phosphinothricin acetyltransferase gene; NOS: A. tumefaciens nopaline synthase poly A; LB: left border.
KJBS-49-180_F1.gif
기상 관측
경북대학교 농업생명과학대학 군위 LMO 격리포장(RDA-가 A-2015-049)에서 해충저항성 Bt 벼(Agb0101)의 유전자 이동 성 평가를 수행하였다. LMO 격리포장에 설치된 기상측정 장치 를 이용하여 개화기간 중 화분의 비산과 수분이 가장 활발한 오전 11를 전후로 4시간 동안 풍속, 풍향, 온도 및 습도를 측정하 였으며, 30분 단위로 기상청에서 실시하는 방식에 따라 기상 자료를 관측하였다.
포장 배치
해충저항성 Bt 벼(Agb0101)의 유전자 이동성을 평가하기 위하여 동, 서, 남, 북, 동북, 동남, 서남, 서북 방향에서 유전자 이동성을 측정할 수 있는 4개의 포장을 조성하였다. 해충저항성 Bt 벼(Agb0101)를 포장의 중심에 1.5×1.5m 크기로 배치하고 해충저항성 Bt 벼(Agb0101)의 상하 방향을 따라서 6m까지 잡초 성벼(R55)와 자포니카 벼품종(낙동벼), 인디카 벼품종(IR36)을 해충저항성 Bt 벼(Agb0101)와 동일한 간격(30×15cm)으로 정 식하였다(Fig. 2).
Fig. 2
An experimental design for the field trial. The GM rice (Agb0101) was planted in white rectangles. Both japonica cultivar (Nakdongbyeo), indica rice (IR36) and weedy rice (R55) were planted following by 8 different directions.
KJBS-49-180_F2.gif
종자 수확
해충저항성 Bt 벼(Agb0101)와 낙동벼, R55, IR36의 개화 진행 상황은 8월 6일부터 약 3주간 조사하였고, 해충저항성 Bt 벼(Agb0101)가 심겨진 마지막 위치를 기준으로 8방향 및 거리(30, 60, 90, 120, 240, 360, 480, 600cm)에 따라 각각 수확하였다. 격리포장 내 건조장에서 건조된 종자를 계수한 후 유전자 이동성을 분석할 때까지 저온실(4℃)에 보관하였다.
유전자 이동성 개체 선별
방향 및 거리별로 낙동벼, R55, IR36로부터 수확한 종자는 혼입을 막기 위해 각각 육묘상자(45×45×5cm)에 고르게 깔고 적절 한 수분을 공급하면서 온실에서 발아시켰다. 3엽기에 0.3% 제초제 (바스타, glufosinate ammonium)를 살포하고 10일 간격으로 2차 와 3차 제초제를 살포하여 제초제저항성을 육안으로 확인하고 저항성 개체를 선발하였다. 선발된 제초제저항성 개체는 분자생물 학적 분석을 위한 시료 채취를 위해 온실에서 재배하였다.
도입유전자의 단백질 발현 검정
선발된 제초제저항성 개체의 mCry1Ac1 유전자의 발현을 확인을 위하여 Immunostrip 검정(lateral strip test)을 실시하였 다. 제초제저항성 개체와 대조구 식물체로부터 유래한 조직으 로부터 단백질을 추출한 후, mCry1Ac1 유전자의 발현을 CryIAc1 ImmunoStrip Test(Agdia, Elkhart, IN)를 이용하여 각각 mCry1Ac1 단백질의 발현 여부를 검정하였다. Immunostrip 검정을 통해 검증된 해충저항성 교잡 개체를 계수 하여 해충저항성 Bt 벼(Agb0101)로부터의 유전자 이동성을 확정하였다.
벼 DNA 분리 및 PCR 검정
Immunostrip 검정을 통해 검증된 해충저항성 교잡개체로부 터 조직시료를 각 0.2 g씩 취하여, 액체질소와 함께 막자 사발에 서 분말화한 후 genomic DNA는 DNeasy Plant kit (Qiagen)를 이용하여 추출하였다. NanoDrop Spectrophotometer ND-1000 (NanoDrop Technologies, Inc, Wilmington, USA)을 이용하여 260/280 nm 값이 1.8-2.0 사이인 추출액을 PCR (Polymerase chain reaction) 실험에 이용하였다. PCR 검정을 위하여 10× PCR buffer 4㎕, dNTP(10mM) 4㎕, f-Taq DNA polymerase 1 unit (Solgent, Korea), 프라이머(Table 1) 각 20μM, 주형 DNA 200pg를 첨가한 후 최종 반응 부피를 40㎕가 되도록 증류수를 첨가하였고, PCR 반응 조건은 1 cycle(95℃, 5분), 35 cycle[95℃(30초), 58℃(30초), 72℃(30초), 1 cycle(72℃, 5분)에 따라 순차적으로 PTC-100 Thermal cycler (MJ Research, USA)를 이용하여 반응시켰다. 각 시료들의 PCR산물 은 1% 한천 겔에서 전기 영동한 후에 UV조사로 확인하였다.
Table 1
Primers list used for Bt-transgenic rice gene flow evaluation.
Table 1
Target Probe Sequences Size (bp)

TpCry1Ac1 C1 5'-CTTCGGCAACGTCAGCAAT-3' 258
C2 5'-GATGGACCAAAGATACCCCAGA-3
Bar B1 5'-CACTACATCGAGACAAGCAC-3 427
B2 5'-TGAAGTCCAGCTGCCAGAAA-3'
Rice genome G1 5'-CCTACATGAGAAATCTTGCTCCTT-3' 342
G2 5'-CCCGAGGATTTTGAGATCAAA-3 '
개화기간 중 기상 조건
Shivrain et al. (2009)는 화분에 의한 유전자 이동성은 식물체 의 화분하(pollen load), 화기형태, 유전적 교잡친화성, 식물체간 의 개화기간 및 기상 조건 등의 환경 요인에 의해 영향을 받는다고 보고하였다(Shivrain et al. 2009). 해충저항성 Bt 벼(Agb0101) 의 화분에 의한 유전자 이동성에 관여하는 기상 조건을 조사하기 위하여 LMO 격리포장 내 설치된 기상 측정 장치를 사용하여 개화 기간 중 온도, 강우량, 풍속 및 풍향을 30분 간격으로 관측하 였다. 잡초성 벼(R55)는 8월 9일부터 개화가 시작되었고, 모품종 인 낙동벼의 개화는 3일 후인 8월 12일부터 시작되었으며, 인디 카 품종인 IR36과 해충저항성 Bt 벼(Agb0101)의 개화는 8월 15일부터 시작되었다. 개화기의 중복이 화분에 의한 인접한 식물 과의 교잡에 주요한 인자이며(Shivrain et al. 2009), 본 실험에서 는 벼의 개화기간이 공시재료간에 근접한 시기에 중복되어 개화 기가 화분에 의한 유전자 이동성 평가에 미치는 영향이 적을 것으로 예상되었다. 관측된 기상자료를 분석한 결과, 개화기간 중 화분의 비산 및 수분이 가장 활발한 오전 9시~12시에는 풍속 0~0.2m/s의 0등급(고요바람)의 상태가 41%를 차지하였으며, 풍속 1등급인 0.3~1.5 m/s의 실바람이 46%, 풍속 1.6~3.3 m/s의 2등급인 남실바람이 8%, 풍속 3.4~5.4 m/s의 3등급인 산들바람 이 5%이었으며, 풍향은 주로 남동풍이 불었다(Fig. 3). 이는 이전에 보고된 경북대 LMO 격리포장의 기상자료와도 유사한 경향을 보였으며(Bae et al. 2013, Oh et al. 2014), 개화기간 동안 총 강수량은 99.2mm였으며, 소폭의 비가 내린 8월 19일부 터 8월 25일까지 기온 저하(22~24℃)가 나타났으며, 이 기간을 제외하고는 온도(26~31℃)와 상대 습도도 80% 이상으로 예년 과 유사한 기상 조건을 나타냈으나, 개화성기(8월 18일)를 전후 하여 내린 비와 기온 저하로 인한 저온다습한 환경이 화분에 의한 유전자 이동성에 영향을 미칠 것으로 사료되었다.
Fig. 3
Variation of meteorological data during flowering. Windrose plot from wind speed and direction (A) and average temperature and rainfall (B).
KJBS-49-180_F3.gif
해충저항성 Bt 벼(Agb0101)의 유전자 이동성
8방향 8단계 거리별로 조성된 포장에서 재배된 모품종인 낙동 벼와 잡초성벼인 R55, 인디카 품종인 IR36로부터 각각 729,917 립, 230,635립, 596,318립의 종자를 수확하였으며, 종자들을 육묘상자에 각 품종별로 파종하여 온실에서 발아시켰다. 선발 마커로 도입된 bar유전자에 의한 제초제인 바스타(glufosinate ammonium) 저항성을 확인하기 위하여 발아된 개체들을 3엽기 까지 생육시킨 후, 제초제(바스타, 0.3%)를 10일 간격으로 3회 살포하여 제초제저항성 여부를 육안으로 판별하였다. 각 품종별로 제초제 저항성으로 확인된 교잡된 개체들로부터 단백질을 추출하 고 해충저항성 Bt 벼 (Agb0101)의 목적 단백질인 mCry1Ac1의 발현 여부를 immunostrip 검정하였다(Fig. 4, Fig. 5, Fig. 6). 제초제 저항성 선별과mCry1Ac1 immunostrip 검정 결과, 모품 종인 낙동벼로부터 제초제 및 해충 저항성을 보이는 20개의 교잡개체를 확인하였으며, 교잡율은 0.0027%로 분석되었다. 해충저항성을 나타내는 낙동벼 교잡개체는 해충저항성 Bt 벼 (Agb0101)에 근접하게 심겨진 동쪽방향(30 cm)에서 1개체, 서쪽 방향(60cm)에서 1개체, 남쪽방향(30 cm, 60 cm, 90 cm)에 서 각각 1개체씩, 북쪽방향에서는 30cm에서 1개체, 60 cm에서 2개체가 발견되었으며, 북서방향에서는 30 cm에서 3개, 60, 90 cm에서 각각 1개체씩 검출되었고, 남서방향에서는 30 cm에서 3개, 120 cm에서 1개체, 남동방향(30 cm)에서 1개체, 북동방향(30, 90 cm)에서 각각 1개체씩 교잡 개체가 검출되었다(Table 2). 잡초 성벼인 R55벼에서의 제초제 및 해충 저항성을 보이는 교잡 개체 는 4개가 발견되었고, 총 파종 종자수에서 교잡 개체 비율(유전자 이동성 비율)은 0.0017%로 분석되었다. 검출된 개체의 방향은 서쪽방향(30, 90 cm)에서 각각 1개체씩, 북서방향(90 cm)에서 1 개체, 북동방향(30 cm)에서 1 개체가 검출되었다(Table 3). 인디카 품종인 IR36벼에서 제초제 및 해충 저항성을 보이는 개체는 3개였으며, 유전자 교잡율은 0.0005%로 확인하였다. 이동된 방향은 동쪽(30 cm)에서 1 개체, 서쪽(30, 90 cm)에서 각각 1개체씩 검출되었다(Table 4). 화분 수용체로 이용된 세 품종인 낙동벼, R55, IR36에서 발견된 총 27개의 교잡개체에서 해충저항성 Bt 벼(Agb0101)에 근접한 1 m이내에서 교잡 개체 의 96.2%가 발견되었다. 동일한 LMO 격리포장에서 연구된 비타민A 강화벼(Bae et al. 2013)와 모품종인 낙동벼에 대한 유전자 이동성 평가에서 제초제 선발에 의한 교잡율은 2009년도 실험 결과에서는 주로 북-서방향 30 cm에서 8.4%(42개체)로 높게 나타났으나, 2010년도 실험 결과에서는 남쪽 30 cm 에서 0.8%(4개체), 동쪽 30 cm에서 0.6%(3개체), 남동쪽 30 cm에서 2.2%(11개체)로 2009년에 비해 낮은 교잡율을 보였다. 비타민A 강화벼 유전자 이동성 실험의 개화성기 기간에 관측된 기상 조건은 2009년도 재배한 벼의 개화기 동안의 바람 방향은 남동풍 (25%), 동남동풍(17.5%), 북서풍(12%) 순서로 불었으며, 최대 풍속은 남동풍 4.9 m/s, 동남동풍과 북서풍은 4 m/s 으로 관측되 었으며, 2010년도의 기상 조건에서 바람의 방향은 남동풍 (21.9%), 남남동풍(17.9%), 동남동풍(10.2%) 순서로 불었으며, 최대 풍속은 남동풍과 남남동은 6.2 m/s, 동남동풍은 5.7 m/s 으로 관측되었다(Bae et al. 2013). 병저항성(OsCK1) 유전자변 형 벼(Oh et al. 2014)의 모품종인 낙동벼에 대한 유전자 이동성 평가에서도 동쪽 30cm에서 0.015%(1개체)와 남쪽 30 cm, 60 cm에서 각각 0.018% (1개체), 0.013% (1개체)로 유전자 이동 빈도는 높지 않았다(Oh et al. 2014). 병저항성 유전자변형벼의 유전자 이동성 실험 기간 동안 격리 포장내의 기상 조건은 개화기 간 중 풍향은 주로 남동풍이 불었으며, 풍속은 바람이 불지 않는 고요한 상태가 49% 였으며, 약풍(0.5-2.0 m/s)은 21.3%, 강풍 (5.7 m/s 이상)의 비율은 2% 이었다. 비록 비타민A 강화벼와 병저항성 유전자변형벼의 다른 유전자가 도입된 벼이지만 동일 한 모품종인 낙동벼에 대한 유전자 이동성 분석시에 개화 성기의 포장내 풍향과 풍속에 따른 유전자 이동률의 차이가 보여주는 결과들은 풍향과 풍속이 화분에 의한 유전자 이동에 따른 교잡 개체의 발생 방향과 빈도에 영향을 주는 환경 요인으로 사료되었 다. 또한, 병저항성 유전자변형벼의 유전자 이동성 실험 기간 동안에 기상 관측시에 개화기간 중의 온도는 28~30℃와 습도는 80% 이상으로 관측되었으나, 개화성기(8월 12일)를 전후하여 두 차례 많은 비가 내린 많은 비로 저온 다습한 환경이 화분에 의한 유전자 이동성에 영향을 주었을 것으로 보여진다(Oh et al. 2014). 이는 동일한 실험 환경에서 잡초성벼(R55)에 대한 유전자 이동성 실험 결과에서도 비타민A 강화벼는 북쪽 30cm에 서 0.4% (2개체), 서쪽 30cm에서 0.4% (2개체), 남서쪽 30cm에 서 1.2% (6개체) 등의 교잡율이 보였으나(Bae et al. 2013), 병저항성 유전자변형 벼에 대한 잡초성벼(R55)의 유전자 이동성 평가에서는 교잡 개체가 검출되지 않았다(Oh et al. 2014). 이와 같은 실험 결과들은 개화기간 중에 풍속, 풍향 뿐만 아니라 저온 다습한 기상 조건이 화분에 의한 유전자 이동에 영향을 미치는 것으로 사료되었다. 이는 벼 유전자 이동성의 환경 요인 중에서 개화시인 오전10시에서 오후 1시까지의 상대습도와 온도는 벼 의 화분과 화기 상태의 변화를 야기하여 화분에 의한 교잡에 영향을 초래할 수 있으며(Shivrain et al. 2009), 따라서, 해충저 항성 Bt 벼(Agb0101) 개화성기의 저온화와 강우에 의한 화분 수용체인 낙동벼, 잡초성벼(R55)로의 화분 이동을 제한시킴으 로써 교잡율이 비타민 A강화벼에 비해 상대적으로 감소하는 경향을 보였으나, 병저항성(OsCK1) 유전자변형 벼에 비해 다소 높은 경향을 보였다. 국내에서 광범위한 지역에 분포하는 잡초성 벼는 벼 재배지 수확량의 22.1%까지 손실을 유발하여 전국적으 로 매년 5~10%의 수확량 감소를 초래되며(Chen et al. 2004), 비형질전환 재배품종과 잡초성벼 간의 교잡률은 실험 조건에 따라 약 0.04~2%로 발생한다고 보고되었다(Messeguer et al. 2004). 이는 제초제 저항성 또는 해충저항성 특성을 지닌 유전자 변형 벼들에 의한 잡초성벼 및 재배종의 유전자 이동성이 발생할 경우 슈퍼잡초 발생 및 재배 품종의 유전자 오염 확산 등 벼 경작지의 생태계에 영향을 미칠 수도 있으나, 본 실험에 공시 재료로 이용된 해충저항성 Bt 벼(Agb0101)는 기존에 보고된 비타민A 강화벼와 병저항성 유전자변형벼에 비해 유사하거나 낮은 교잡율이 보였기에 유전자 이동성에 의한 농업생태계의 위험은 크지 않을 것으로 추정된다. 그러나, 벼 재배지 내의 유전자 이동성에 대한 농업생태계의 위해성 평가는 벼의 개화성 기 중의 재배 지역의 기상 조건들과 벼 품종간의 개화기 차이 등의 변화에 따른 다년간의 평가와 과학적인 데이터 분석을 통한 체계적인 접근이 요구되며, 이를 기반으로 벼의 유전자 이동성 감축 기술 개발과 유전자 이동성의 모델링 통한 사전 예측과 관련된 연구가 수행되어야 할 것이다.
Fig. 4
Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and japonica non-GM rice by using immunostrip. C: non-GM rice (Nakdongbyeo), GM: Bt transgenic rice (Agb0101), NHY 1~20: Nakdongbyeo × Agb0101 hybrids.
KJBS-49-180_F4.gif
Fig. 5
Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and weedy rice by using immunostrip. C: weedy rice (R55), GM: Bt transgenic rice (Agb0101), RHY 1~4: R55 × Agb0101 hybrids.
KJBS-49-180_F5.gif
Fig. 6
Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and indica non-GM rice by using immunostrip. C: non-GM rice (IR36), GM: Bt transgenic rice (Agb0101), IHY 1~3: IR36 × Agb0101 hybrids.
KJBS-49-180_F6.gif
Table 2
Out-crossing rate (%) of japonica non-GM rice (Nakdongbyeo) at different distance and direction from GM pollen donor.
Table 2
Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0.008 0 0.006 0.007 0.023 0.018 0.008 0.008
(1/11,555)Z (0/11,953) (1/14,593) (1/12,967) (3/12,669) (3/16,287) (1/11,155) (1/12,269)
60 0 0.012 0.010 0.017 0.008 0 0 0
(0/9,000) (1/7,838) (1/9,888) (2/11,427) (1/12,194) (0/13,134) (0/8,435) (0/10,844)
90 0 0 0.008 0 0.009 0 0 0.008
(0/11,068) (0/9,858) (1/11,957) (0/9,975) (1/10,667) (0/12,435) (0/10,913) (1/11,683)
120 0 0 0 0 0 0.007 0 0
(0/11,413) (0/9,447) (0/12,653) (0/11,010) (0/13,608) (1/12,716) (0/12,920) (0/10,016)
240 0 0 0 0 0 0 0 0
(0/10,925) (0/9,007) (0/12,730) (0/10,143) (0/11,747) (0/9,367) (0/12,101) (0/13,038)
360 0 0 0 0 0 0 0 0
(0/10,968) (0/9,252) (0/12,523) (0/9,884) (0/9,705) (0/7,925) (0/12,231) (0/15,383)
480 0 0 0 0 0 0 0 0
(0/9,083) (0/11,352) (0/7,934) (0/10,080) (0/11,156) (0/9,541) (0/13,754) (0/15,729)
600 0 0 0 0 0 0 0 0
(0/11,287) (0/8,853) (0/11,484) (0/11,998) (0/13,506) (0/11,947) (0/13,720) (0/13,018)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Table 3
Out-crossing rate (%) of weedy rice (R55) at different distance and direction from GM pollen donor.
Table 3
Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0 (0/4,066)Z 0.030 (1/3,257) 0 (0/6,610) 0 (0/4,168) 0 (0/6,866) 0 (0/5,366) 0 (0/3,123) 0.019 (1/5,191)
60 0 (0/3,808) 0 (0/3,614) 0 (0/3,303) 0 (0/4,528) 0 (0/3,833) 0 (0/2,875) 0 (0/2,005) 0 (0/2,039)
90 0 (0/3,560) 0.035 (1/2,843) 0 (0/6,037) 0 (0/4,648) 0.028 (1/3,521) 0 (0/593) 0 (0/4,214) 0 (0/1,896)
120 0 (0/2,897) 0 (0/4,045) 0 (0/5,178) 0 (0/4,371) 0 (0/2,394) 0 (0/837) 0 (0/3,776) 0 (0/2,976)
240 0 (0/3,362) 0 (0/4,191) 0 (0/4,938) 0 (0/4,741) 0 (0/1,650) 0 (0/2,583) 0 (0/3,206) 0 (0/4,439)
360 0 (0/4,223) 0 (0/5,253) 0 (0/3,669) 0 (0/1,478) 0 (0/2,691) 0 (0/1,827) 0 (0/4,062) 0 (0/3,505)
480 0 (0/4,081) 0 (0/4,740) 0 (0/3,570) 0 (0/3,236) 0 (0/1,937) 0 (0/2,955) 0 (0/3,698) 0 (0/2,304)
600 0 (0/3,641) 0 (0/2,806) 0 (0/4,320) 0 (0/3,833) 0 (0/2,941) 0 (0/5,111) 0 (0/5,533) 0 (0/1,674)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Table 4
Out-crossing rate (%) of indica non-GM rice (IR36) at different distance and direction from GM pollen donor.
Table 4
Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0.008 (1/11,740)Z 0.008 (1/11,456) 0 (0/12,092) 0 (0/12,245) 0 (0/11,636) 0 (0/7,648) 0 (0/8,726) 0 (0/11,236)
60 0 (0/10,354) 0 (0/8,864) 0 (0/7,843) 0 (0/11,647) 0 (0/8,748) 0 (0/3,533) 0 (0/7,370) 0 (0/6,601)
90 0 (0/10,311) 0.009 (1/10,510) 0 (0/10,109) 0 (0/8,265) 0 (0/9,210) 0 (0/4,272) 0 (0/7,033) 0 (0/6,654)
120 0 (0/8,038) 0 (0/10,597) 0 (0/12,011) 0 (0/9,049) 0 (0/7,725) 0 (0/6,761) 0 (0/7,152) 0 (0/8,660)
240 0 (0/10,931) 0 (0/12,091) 0 (0/8,946) 0 (0/10,512) 0 (0/8,948) 0 (0/6,816) 0 (0/7,501) 0 (0/8,663)
360 0 (0/11,522) 0 (0/10,605) 0 (0/8,971) 0 (0/12,362) 0 (0/11,533) 0 (0/6,954) 0 (0/9,334) 0 (0/9,265)
480 0 (0/9,047) 0 (0/11,254) 0 (0/10,037) 0 (0/10,693) 0 (0/9,326) 0 (0/9,863) 0 (0/5,714) 0 (0/8,381)
600 0 (0/7,855) 0 (0/11,324) 0 (0/10,469) 0 (0/12,084) 0 (0/10,295) 0 (0/8,583) 0 (0/9,201) 0 (0/9,141)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

유전자 이동성 개체의 PCR 검정
제초제(바스타) 살포를 통해 제초제 저항성을 보이는 유전자 이동성 개체에 대한 분자생물학적 검정을 위해 선별된 식물 재료들로부터 DNA를 추출하고 PCR 분석을 수행하였다. 추출 된 DNA를 주형으로 해충저항성 Bt 벼(Agb0101)에 도입된 목적유전자(TpCry1Ac1)와 제초제저항성 유전자(Bar)에 특이적 인 프라이머를 사용하여 밴드의 증폭을 확인하였다(Table 1). 모품종인 낙동벼는 해충저항성 유전자 (TpCry1Ac1)와 제초제저 항성 유전자(Bar)가 도입되지 않았기 때문에 PCR 밴드가 검출 되지 않았으며, Agb0101와 교잡된 20개의 개체들에서는 제초 제 선별 및 mCry1Ac1 immunostrip 검정 결과와 일치되는 TpCry1Ac1 유전자와 Bar 유전자의 밴드가 검출되어 교잡된 개체의 도입된 유전자가 Agb0101로부터 유래된 것을 확인하였 다. 또한, 고정된 해충저항성 Bt 벼(Agb0101)와는 달리 교잡된 개체(F1 세대)는 모계(낙동벼)와 부계(Agb0101)의 양쪽에서 유래한 게놈이 조합되었기 때문에 Agb0101와 교잡된 개체들에 서 rice genome 유전자(Agb0101의 T-DNA가 도입된 주변 염기 서열)를 PCR을 분석한 결과에서 낙동벼와 동일한 크기의 밴드 가 검출됨을 확인하였다(Fig. 7). 잡초성벼(R55)의 교잡 개체 4개와 인디카벼(IR36)의 교잡개체 3개에 대한 PCR 검정 결과에 서도 TpCry1Ac1 유전자와 Bar 유전자의 밴드가 검출되어 교잡 된 개체가 Agb0101로부터 유래된 것임을 확인하였으며, rice genome 유전자의 PCR 분석에서도 낙동벼 교잡개체들과 동일 한 결과를 확인하였다(Fig. 8, 9). rice genome 유전자 PCR 수행 결과에서도 Agb0101벼에 도입유전자(T-DNA)의 크기가 약 9.5 kb로 삽입되어 있어 수행한 PCR 반응 조건으로 DNA 증폭하기 어려운 크기로 인해 Agb0101벼의 rice genome 유전 자의 밴드가 검출이 되지 않았다. 전 세계적으로 유전자변형 (Genetically Modified, GM)작물은 식량위기 해결과 농약 사용 량 절감에 따른 경제적 이익뿐만 아니라 환경보호 등의 효과에도 불구하고 농업환경 및 식품 안전성 때문에 현재까지 논쟁 중이다 (Bae et al. 2013). GM작물의 생산, 이용 등에 반대하는 측은 재배에 따른 농업 환경 변화와 섭식에 따른 인간에 미치는 연구가 불충분하여 안전성이 검증되지 않았다고 우려한다. 현재까지 국내에서는 GM벼를 포함한 어떠한 생명공학 기술을 이용한 GM작물이 상업적으로 재배되고 있지 않지만, GM작물로부터 유전자 이동에 의한 재배종의 유전자 오염, 슈퍼 잡초 발생 및 종 다양성 영향 등의 문제를 발생할 수 있으며, 유전자 이동성의 비율이 높지 않더라도 주변 농업 환경에 충분히 영향을 줄 수 있으므로, GM작물의 재배에 의한 재배⋅근연종으로의 유전자 이동성을 최소화하는 안전관리 방법과 대응 매뉴얼의 개발이 반드시 필요하다. 이는 작물 재배 지역의 기상 환경 요소와 화분 매개충의 발생 유무 및 유전자변형작물과 재배⋅근연종의 생리 적, 형태학적 특성을 다변화된 변수가 포함된 유전자 이동 모델링 구축과 개발이 선행되어야 할 것이다.
Fig. 7
Detection of hybrids between GM rice and japonica non-GM rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar gene and rice genome for hybrid rice. M: DNA marker, C: non-GM rice (Nakdongbyeo), GM: Bt transgenic rice (Agb0101), NHY 1~20: Nakdongbyeo × Agb0101 hybrids.
KJBS-49-180_F7.gif
Fig. 8
Detection of hybrids between GM rice and weedy rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar gene and rice genome for hybrid rice. M: DNA marker, C: weedy rice (R55), GM: Bt transgenic rice (Agb0101), RHY 1~4: R55 × Agb0101 hybrids.
KJBS-49-180_F8.gif
Fig. 9
Detection of hybrids between GM rice and indica non-GM rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar1111111111 gene and rice genome for hybrid rice. M: DNA marker, C: non-GM rice (IR36), GM: Bt transgenic rice (Agb0101), IHY 1~3: IR36 × Agb0101 hybrids.
KJBS-49-180_F9.gif
현재까지 국내에서는 생명공학작물이 상업적인 재배가 되고 있지 않으나 생명공학작물의 환경 방출을 위해서는 환경위해성 평가가 필수적 수행되어야 한다. 본 연구에서는 해충저항성 Bt 벼(Agb0101)로부터 모품종인 낙동벼와 잡초성벼인 R55 및 인 디카벼인 IR36 로의 화분 매개에 의한 유전자 이동성을 평가하였 다. 낙동벼로부터 729,917 립의 종자를 얻었으며, 잡초성벼 (R55)로부터는 230,635 립의 종자와 인디카벼(IR36)에서는 596,318 립의 종자를 수확하였다. 교잡개체는 3회의 제초제 살포를 수행하여 제초제 저항성 개체 선별과 Cry1Ac1 immunostrip 검정으로 확인하였으며, 해충저항성 Bt 벼 (Agb0101)에 특이적인 프라이머를 이용한 분자생물학적 방법 을 통해 유전자 이동성 여부를 최종적으로 검증하였다. 총 파종된 종자수에 대한 교잡율은 낙동벼에서는 0.0027%, R55는 0.0017%, IR36은 0.0005%로 나타났으며, 모든 교잡개체들은 해충저항성 Bt 벼(Agb0101)에 근접한 1.2m 내에서 발견되었다. 화분 매개에 의한 해충저항성 Bt 벼(Agb0101)의 유전자 이동 특성은 기존에 연구된 결과들과 유사한 경향으로 보였으며, 벼의 개화기간 중 온도와 강우량 등 기상 조건이 화분에 의한 교잡을 결정하는데 중요한 요인으로 작용하였다. 이에 재배 지역의 기상 환경과 개화시기 중복 여부 등을 유전자변형 벼에 의한 일반 재배품종 및 야생(잡초성)벼로의 유전자 이동에 따른 저감 기술 개발과 안전관리 기준 작성에서 주요 영향 인자들로 고려되어야 할 것이다.
본 연구는 농촌진흥청 국립농업과학원 농업과학기술 연구개 발사업(과제번호: PJ01186701)의 지원에 의해 이루어진 것임.
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Assessment of gene flow from insect-resistant genetically modified rice (Agb0101) to non-GM rice
Korean. J. Breed. Sci.. 2017;49(3):180-189.   Published online September 1, 2017
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Assessment of gene flow from insect-resistant genetically modified rice (Agb0101) to non-GM rice
Korean. J. Breed. Sci.. 2017;49(3):180-189.   Published online September 1, 2017
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Assessment of gene flow from insect-resistant genetically modified rice (Agb0101) to non-GM rice
Image Image Image Image Image Image Image Image Image
Fig. 1 Schematic representation of the inserted vector used for rice transformation. The arrows indicate primer (Table 1) sites and the direction of synthesis. RB: right border; MAR: matrix attachment region; rbsS: rice rbcS promoter; TP: rice rbcS transit peptide; mCry1Ac1: B. thuringiensis mCry1Ac1 gene; PinII: potato proteinase inhibitor II gene; 35S: CaMV 35S promoter; PAT: phosphinothricin acetyltransferase gene; NOS: A. tumefaciens nopaline synthase poly A; LB: left border.
Fig. 2 An experimental design for the field trial. The GM rice (Agb0101) was planted in white rectangles. Both japonica cultivar (Nakdongbyeo), indica rice (IR36) and weedy rice (R55) were planted following by 8 different directions.
Fig. 3 Variation of meteorological data during flowering. Windrose plot from wind speed and direction (A) and average temperature and rainfall (B).
Fig. 4 Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and japonica non-GM rice by using immunostrip. C: non-GM rice (Nakdongbyeo), GM: Bt transgenic rice (Agb0101), NHY 1~20: Nakdongbyeo × Agb0101 hybrids.
Fig. 5 Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and weedy rice by using immunostrip. C: weedy rice (R55), GM: Bt transgenic rice (Agb0101), RHY 1~4: R55 × Agb0101 hybrids.
Fig. 6 Confirmation of Cry1Ac1 protein expression for hybrids between GM rice and indica non-GM rice by using immunostrip. C: non-GM rice (IR36), GM: Bt transgenic rice (Agb0101), IHY 1~3: IR36 × Agb0101 hybrids.
Fig. 7 Detection of hybrids between GM rice and japonica non-GM rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar gene and rice genome for hybrid rice. M: DNA marker, C: non-GM rice (Nakdongbyeo), GM: Bt transgenic rice (Agb0101), NHY 1~20: Nakdongbyeo × Agb0101 hybrids.
Fig. 8 Detection of hybrids between GM rice and weedy rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar gene and rice genome for hybrid rice. M: DNA marker, C: weedy rice (R55), GM: Bt transgenic rice (Agb0101), RHY 1~4: R55 × Agb0101 hybrids.
Fig. 9 Detection of hybrids between GM rice and indica non-GM rice. Three pairs of primers used to detect TpCry1Ac1 gene, bar1111111111 gene and rice genome for hybrid rice. M: DNA marker, C: non-GM rice (IR36), GM: Bt transgenic rice (Agb0101), IHY 1~3: IR36 × Agb0101 hybrids.
Assessment of gene flow from insect-resistant genetically modified rice (Agb0101) to non-GM rice

Primers list used for Bt-transgenic rice gene flow evaluation.

Target Probe Sequences Size (bp)

TpCry1Ac1 C1 5'-CTTCGGCAACGTCAGCAAT-3' 258
C2 5'-GATGGACCAAAGATACCCCAGA-3
Bar B1 5'-CACTACATCGAGACAAGCAC-3 427
B2 5'-TGAAGTCCAGCTGCCAGAAA-3'
Rice genome G1 5'-CCTACATGAGAAATCTTGCTCCTT-3' 342
G2 5'-CCCGAGGATTTTGAGATCAAA-3 '

Out-crossing rate (%) of japonica non-GM rice (Nakdongbyeo) at different distance and direction from GM pollen donor.

Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0.008 0 0.006 0.007 0.023 0.018 0.008 0.008
(1/11,555)Z (0/11,953) (1/14,593) (1/12,967) (3/12,669) (3/16,287) (1/11,155) (1/12,269)
60 0 0.012 0.010 0.017 0.008 0 0 0
(0/9,000) (1/7,838) (1/9,888) (2/11,427) (1/12,194) (0/13,134) (0/8,435) (0/10,844)
90 0 0 0.008 0 0.009 0 0 0.008
(0/11,068) (0/9,858) (1/11,957) (0/9,975) (1/10,667) (0/12,435) (0/10,913) (1/11,683)
120 0 0 0 0 0 0.007 0 0
(0/11,413) (0/9,447) (0/12,653) (0/11,010) (0/13,608) (1/12,716) (0/12,920) (0/10,016)
240 0 0 0 0 0 0 0 0
(0/10,925) (0/9,007) (0/12,730) (0/10,143) (0/11,747) (0/9,367) (0/12,101) (0/13,038)
360 0 0 0 0 0 0 0 0
(0/10,968) (0/9,252) (0/12,523) (0/9,884) (0/9,705) (0/7,925) (0/12,231) (0/15,383)
480 0 0 0 0 0 0 0 0
(0/9,083) (0/11,352) (0/7,934) (0/10,080) (0/11,156) (0/9,541) (0/13,754) (0/15,729)
600 0 0 0 0 0 0 0 0
(0/11,287) (0/8,853) (0/11,484) (0/11,998) (0/13,506) (0/11,947) (0/13,720) (0/13,018)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Out-crossing rate (%) of weedy rice (R55) at different distance and direction from GM pollen donor.

Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0 (0/4,066)Z 0.030 (1/3,257) 0 (0/6,610) 0 (0/4,168) 0 (0/6,866) 0 (0/5,366) 0 (0/3,123) 0.019 (1/5,191)
60 0 (0/3,808) 0 (0/3,614) 0 (0/3,303) 0 (0/4,528) 0 (0/3,833) 0 (0/2,875) 0 (0/2,005) 0 (0/2,039)
90 0 (0/3,560) 0.035 (1/2,843) 0 (0/6,037) 0 (0/4,648) 0.028 (1/3,521) 0 (0/593) 0 (0/4,214) 0 (0/1,896)
120 0 (0/2,897) 0 (0/4,045) 0 (0/5,178) 0 (0/4,371) 0 (0/2,394) 0 (0/837) 0 (0/3,776) 0 (0/2,976)
240 0 (0/3,362) 0 (0/4,191) 0 (0/4,938) 0 (0/4,741) 0 (0/1,650) 0 (0/2,583) 0 (0/3,206) 0 (0/4,439)
360 0 (0/4,223) 0 (0/5,253) 0 (0/3,669) 0 (0/1,478) 0 (0/2,691) 0 (0/1,827) 0 (0/4,062) 0 (0/3,505)
480 0 (0/4,081) 0 (0/4,740) 0 (0/3,570) 0 (0/3,236) 0 (0/1,937) 0 (0/2,955) 0 (0/3,698) 0 (0/2,304)
600 0 (0/3,641) 0 (0/2,806) 0 (0/4,320) 0 (0/3,833) 0 (0/2,941) 0 (0/5,111) 0 (0/5,533) 0 (0/1,674)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Out-crossing rate (%) of indica non-GM rice (IR36) at different distance and direction from GM pollen donor.

Distance (cm) East West South North Northwest Southwest Southeast Northeast

30 0.008 (1/11,740)Z 0.008 (1/11,456) 0 (0/12,092) 0 (0/12,245) 0 (0/11,636) 0 (0/7,648) 0 (0/8,726) 0 (0/11,236)
60 0 (0/10,354) 0 (0/8,864) 0 (0/7,843) 0 (0/11,647) 0 (0/8,748) 0 (0/3,533) 0 (0/7,370) 0 (0/6,601)
90 0 (0/10,311) 0.009 (1/10,510) 0 (0/10,109) 0 (0/8,265) 0 (0/9,210) 0 (0/4,272) 0 (0/7,033) 0 (0/6,654)
120 0 (0/8,038) 0 (0/10,597) 0 (0/12,011) 0 (0/9,049) 0 (0/7,725) 0 (0/6,761) 0 (0/7,152) 0 (0/8,660)
240 0 (0/10,931) 0 (0/12,091) 0 (0/8,946) 0 (0/10,512) 0 (0/8,948) 0 (0/6,816) 0 (0/7,501) 0 (0/8,663)
360 0 (0/11,522) 0 (0/10,605) 0 (0/8,971) 0 (0/12,362) 0 (0/11,533) 0 (0/6,954) 0 (0/9,334) 0 (0/9,265)
480 0 (0/9,047) 0 (0/11,254) 0 (0/10,037) 0 (0/10,693) 0 (0/9,326) 0 (0/9,863) 0 (0/5,714) 0 (0/8,381)
600 0 (0/7,855) 0 (0/11,324) 0 (0/10,469) 0 (0/12,084) 0 (0/10,295) 0 (0/8,583) 0 (0/9,201) 0 (0/9,141)

zOut-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Table 1 Primers list used for Bt-transgenic rice gene flow evaluation.
Table 2 Out-crossing rate (%) of japonica non-GM rice (Nakdongbyeo) at different distance and direction from GM pollen donor.

Out-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Table 3 Out-crossing rate (%) of weedy rice (R55) at different distance and direction from GM pollen donor.

Out-crossing rate (%) (No. of hybrid plant/No. of sown seed)

Table 4 Out-crossing rate (%) of indica non-GM rice (IR36) at different distance and direction from GM pollen donor.

Out-crossing rate (%) (No. of hybrid plant/No. of sown seed)