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Delta 15 desaturase 유전자 억제에 의해 알파리놀렌산 함량이 낮은 들깨 육성

김경환1,*, 이경렬1, 김정봉2, 이명희3, 이은경1, 김년희1, 이홍석1, 김송림1, 백정호1, 최인찬1, 지현소1

Development of Perilla frutescens with Low Levels of Alpha-Linolenic Acid by Inhibition of a delta 15 desaturase Gene

Kyung-Hwan Kim1,*, Kyeong-Ryeol Lee1, Jung-Bong Kim2, Myoung Hee Lee3, Eungyeong Lee1, Nyunhee Kim1, Hongseok Lee1, Song Lim Kim1, JeongHo Baek1, Inchan Choi1, Hyeonso Ji1
Korean Journal of Breeding Science 2018;50(4):463-471.
Published online: December 1, 2018

농촌진흥청 국립농업과학원 농업생명자원부

농촌진흥청 국립농업과학원 농식품자원부

농촌진흥청 국립식량과학원 남부작물부

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* (E-mail: biopiakim@korea.kr, Tel: +82-63-238-4658, Fax: +82-63-238-4654)
• Received: October 30, 2018   • Revised: October 30, 2018   • Accepted: November 7, 2018

Copyright: © 2018 by the Korean Society of Breeding Science

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  • Perilla is an oilseed crop cultivated in Korea since ancient times. Due to the high α-linolenic acid content in perilla, perilla seed oil can easily become rancid. α-Linolenic acid is synthesized by two enzymes, endoplasmic reticulum-localized Δ15 desaturase (FAD3) and chloroplast-localized Δ15 desaturase (FAD7) in vivo. In order to lower the α-linolenic acid content of the seed oil without disturbing plant growth, we tried to suppress the expression of only the FAD3 gene using RNA interference, whilst maintaining the expression of the FAD7 gene. Seventeen transgenic plants with herbicide (Basta™) resistance were obtained by Agrobacterium-mediated transformation using hypocotyls of perilla plants. The transgenic plants were firstly confirmed by treatment with 0.3% (v/v) Basta™ herbicide, and the expression of FAD3 was measured by Northern blot analysis. The α-linolenic acid content was 10-20%, 30-40%, and 60% in two, seven, and three of the twelve T1 transgenic perilla plants which had enough seeds to be analyzed for fatty acid composition, respectively. Analysis of the fatty acid composition of T2 progeny seeds from T1 plants with the lowest α-linolenic acid content showed that the homozygous lines had 6-10% α-linolenic acid content and the heterozygous lines had 20-26% α-linolenic acid content. It is expected that the reduction in α-linolenic acid content in perilla seed oil will prevent rancidity and can be utilized for the production of high-value functional ingredients such as high γ-linolenic acid.
  • Keywords: FAD3; Δ15 desaturase; α-linolenic acid; RNAi; Perilla
들깨(Perilla frutescens var. frutescens)는 우리나라를 비롯한 동북아 일대에서 고대로부터 재배되어 온 유지작물이다. 종자는 지질51%, 단백질 17%, 탄수화물 20% 정도이며 지질이 대부분을 차지하고 있어 식용유로 많이 사용하고 있으며(Longvah et al. 1991), 잎은 최근 신선채소로 각광받고 있다. 종자 지방산의 조성은 올레산(oleic acid, 18:1Δ9) 14-23%, 리놀레산(linoleic acid, 18:2Δ9,12) 11-16%, 알파리놀렌산(α-linolenic acid, 18:3Δ9,12,15) 54-64%로 필수지방산인 리놀레산과 알파리놀렌산이 대부분을 차지하여 건강에 이로우며 알파리놀렌산은 심혈관계 질환, 암, 염증, 류마티스 등의 질병 예방 효과가 있다고 알려져 있다(Asif 2011, Shin & Kim 1994). 또한 들깨는 아마(linseed), 치아(chia)와 더불어 종자에서 알파리놀렌산의 함량이 가장 높은 유지작물로 최근 오메가3 지방산의 주요 공급원으로 각광받고 있다(Ciftci et al. 2012). 이런 좋은 장점에도 불구하고 가장 높은 비율을 차지하는 알파리놀렌산 때문에 들기름의 산패가 쉽게 일어나 저장이 어려운 단점이 있다. 이에 산패를 방지하기 위해서 들깨에 부족한 리그난(세사미놀, 세사몰린 등) 같은 항산화물질을 추가하거나 들깨에서 알파리놀렌산 합성효소의 발현을 억제하여 알파리놀렌산 함량이 낮은 들깨를 만들 필요성이 있다. 또한 알파리놀렌산의 합성을 억제함으로써 리놀레산을 많이 축적하여 들깨에서 생성되지 않는 감마리놀렌산(γ-linolenic acid, 18:3Δ6,9,12) 같은 건강기능성이 있는 특이 지방산 생산시 기질로 사용할 수 있기 때문에 생명공학적으로 대사경로를 조절하여 알파리놀렌산을 감소시키고 리놀레산 함량을 증가할 필요성이 있다. 알파리놀렌산을 합성하는 효소는 두 종류가 있는데 소포체 막에 저장지질을 구성하며 알파리놀렌산을 합성하는 Δ15 desaturase (Fatty Acid Desaturase 3; FAD3)와 엽록체 막에 존재하며 세포막의 지방산 합성에 관여하는 Δ15 desaturase (FAD7, FAD8)이다. 이 두 효소는 서로간의 아미노산 서열이 매우 유사하나 FAD7의 경우 FAD3에는 없는 엽록체 이동 펩타이드(Chloroplast transit peptide)가 N-terminal에 존재한다(Iba et al. 1993, Lee et al. 2016). FAD8은 저온으로 FAD7의 발현이 저해를 받을 때 오히려 발현이 유도되어 엽록체 내의 불포화도를 조절하는 역할을 하며(Mcconn et al. 1994) 들깨에서는 보고되지 않았다. FAD7 유전자의 발현이 억제되어 영양조직의 엽록체 지질 내에 알파리놀렌산의 비율이 낮아지게 되면 생장에 저해를 입을 수 있기 때문에 종자의 저장지질의 알파리놀렌산 합성과 관계된 FAD3 유전자의 발현을 억제하는 것이 생장 저해를 받지 않고 종자 지방산 내 알파리놀렌산 비율을 줄일 수 있다.
생물체 내의 유전자의 발현을 억제하는 방법에는 여러 가지가 개발되어 사용되고 있다. 먼저 발현을 억제할 유전자를 antisense 방향으로 발현하여 기존 유전자의 mRNA와 결합하게 하여 단백질 합성을 막는 antisense RNA 방법이 있고(Ecker & Davis 1986), 발현을 억제할 유전자를 과발현하여 기존 유전자의 silencing을 유도하는 cosuppression 방법이 있다(Napoli et al. 1990). 또한 유전자의 일부분만을 이용하여 인트론 앞뒤로 sense 및 antisense 방향으로 붙여 mRNA가 hairpin loop를 형성케 하고 이때 만들어진 이중나선 RNA가 Dicer에 의해 21-23 bp의 small interfering RNA (siRNA)로 나눠지면서 기존 유전자의 mRNA에 붙고 RNA-induced Silencing Complex (RISC)가 이를 분해함으로써 유전자 발현을 억제하는 RNA 간섭(RNA interference, RNAi) 방법 등이 있다(Fire et al. 1998).
이 연구에서는 RNAi 방법을 이용하여 FAD7의 발현은 억제하지 않으면서 FAD3 유전자의 발현만 억제하기 위한 유전자 운반체를 제작하였고 이를 들깨에 도입하여 형질전환하였으며 종자지방산 분석 결과 비형질전환 들깨에 비해 알파리놀렌산 함량이 매우 낮은 형질전환 들깨를 개발하고자 하였다.
형질전환 운반체 작성
GenBank (https://www.ncbi.nlm.nih.gov/genbank/)에 등록된 들깨의 소포체 Δ15 desaturase (FAD3) 유전자(Genbank accession No. AF213482)와 엽록체 Δ15 desaturase (FAD7) 유전자(Genbank accession No. KC442313)의 아미노산 서열을 비교하여 상동성이 가장 낮은 부분에 해당하는 193 bp 크기의 염기서열을 RNAi 운반체 제작에 이용하였다. 운반체 제작에 사용한 유전자 단편은 Gateway 클로닝 방법을 위해 특이적 프라이머와 P1-P2 PCR을 통해 증폭하였다. 사용된 Gateway 클로닝 방법 및 PCR 조건은 제조사의 방법을 따랐다. 이 실험에 사용된 특이적 프라이머의 염기서열은 다음과 같다. P1 PCR primer: Fad3-B1, 5’-AAAAAGCAGGCTGAGAGGGGCTTGATAGT-3’; Fad3-B2, 5’-AGAAAGCTGGGTGGTACCAAGGGAGTTTC-3’; P2 PCR primer: attB1, 5’-GGGGACAAGTTTGTACAAAAAAGCAGGCT-3’; attB2, 5’-GGGGACCACTTTGTACAAGAAAGCTGGGT-3’. 증폭된 유전자 단편은 Gateway BP clonase (Invitrogen, USA)와 pDONR201 (Invitrogen, USA)를 섞고 25°C에서 2시간 이상을 두어 재조합시켜 pENTR-FAD3-193 유전자 운반체가 제작되었다. 최종 식물형질전환 운반체를 제작하기 위해 목적유전자를 pB7GWIWG2(II) 운반체(Karimi et al. 2002), pENTR-FAD3-193운반체 및 Gateway LR clonase (Invitrogen, USA)와 혼합한 후 25°C에서 4시간 이상 재조합 반응을 거쳐 최종 운반체인 pF3SiA가 완성되었다. 제작된 형질전환용 운반체들은 액체질소를 이용한 동결-해동법(freeze-thaw method, Holster et al. 1978)으로 Agrobacterium tumefaciens 균주 GV3101에 형질전환 시킨 후, PCR을 이용하여 아그로박테리움으로 도입을 확인하였다.
들깨 형질전환
들깨 형질전환은 배축을 이용하여 직접 신초를 형성하고 제초제(phosphinothricin)를 이용하여 선발하는 기존의 방법 에 따라 실시하였다(Kim et al. 2004). 들깨 종자는 70% (v/v) 에탄올에 1분간 침지한 후 50% (v/v) 상업용 표백제에서 20분간 살균하고 멸균수로 3-5회 수세한 다음 물기를 제거하여 MS 기본 배지(Murashige & Skoog 1962)가 든 petridish에 파종 후 6-8일의 배축을 이용하여 형질전환 재료로 사용하였다. 배축을 0.5-1 cm 크기로 절단한 후 pF3SiA 운반체를 가진 Agrobacterium tumefaciens GV3101과 3일간 공동배양 후 cefotaxime 250 mg/L가 첨가된 MS 액체배지(MS salt+vitamin, pH 5.6)로 세척하였다. 여분의 수분을 제거한 절편을 신초 유도배지[MS salt+vitamin, 3% (w/v) sucrose, 3 mg/L BA, 0.1 mg/L NAA, 0.2% gelrite, 0.2% phyto agar, 1.2 mg/L PPT, 500 mg/L carbenicillin, pH 5.6]에 옮겨서 형질전환된 배축에서만 신초가 유도되도록 하였다. 유도된 신초 부분만을 절단하여 신초 신장배지(MS salt+vitamin, 3 mg/L BA, 3% sucrose, 0.2% gelrite, 0.2% phyto agar, 2 mg/L PPT, 500 mg/L carbenicillin, pH 5.6)에서 2달 계대한 후 뿌리 유도배지(1/2 MS salt+vitamin, 1.5% sucrose, 0.2% gelrite, 0.2% phyto agar, 500 mg/L carbenicillin, pH 5.6)에서 뿌리 생성하였다. 뿌리가 생성된 식물체는 순화과정을 거쳐 온실에서 재배하였다.
형질전환 들깨의 유전자발현 확인
재분화된 들깨가 성장하여 본엽이 5-8개의 나왔을 때 0.3% (v/v) 바스타 제초제를 살포하여 형질전환여부를 우선 판별하였다. 바스타 제초체에서 생존한 개체를 대상으로 잎과 미숙 종자에서 유전자의 발현을 확인하였다. 어린 잎과 수정 후 15일 된 미숙종자를 채취하여 Smith 등(1988)의 방법에 따라 total RNA를 분리하고 193 bp 크기의 FAD3 유전자 단편을 탐침(probe)으로 사용하여 Northern blot으로 유전자 발현을 확인하였다(Alwine et al. 1977). 온실에서 재배하여 수확한 형질전환 종자는 분리비 검정과 지방산 분석의 재료로 사용되었다.
형질전환 들깨종자의 지방산 분석
형질전환들깨의 지방산을 분석하기 위하여 0.1 g의 들깨 종자를 대상으로 막자와 막자사발을 이용하여 분쇄하고 1 ml의 내부표준물질(pentadecanoic acids in MeOH, 1000 ppm)과 1 ml의 추출용매(chloroform:MeOH=2:1)를 첨가한 후 4 ml test tube에 넣어 지방산을 용출하고 5 ml의 0.58% NaCl 용액을 넣고 10분간 sonication하였다. 추출액을 원심분리한 후 상층액을 버리고 회전농축기(rotary evaporator)를 이용하여 시료를 농축한 후 농축한 시료에 0.5 ml의 toluene과 2 ml의 0.5 N NaOH in MeOH를 첨가하여 끓는 물에 5분간 반응시켜 지방산을 용출하였다. 이후 2.5 ml의 14% (w/v) BF3 in MeOH를 첨가한 후 끓는 물에서 5분간 반응시켜 지방산을 methylation 시켜 methyl ester로 만든 후 10 ml의 petroleum ether와 15 ml의 3차 증류수를 넣고 잘 섞은 후 10분간 sonication하였다. 반응액을 Na2SO4를 이용하여 상등액을 filtration한 후에 농축하고 1 ml petroleum ether를 넣은 후 HP5890 (Agilent, USA) 가스크로마토그래프를 이용하여 분석하였다. 지방산 분석을 위한 검출기로는 불꽃이온화 검출기(flame ionization detector, FID)를 사용하였으며 컬럼은 DB-WAX (30 m, 0.25 mm, 0.25 μm, Agilent, USA)를 사용하였다. 분석조건은 다음과 같다. 불꽃점화 및 연소를 위한 공기는 370 ml/min로 공급되었으며 수소가스는 47 ml/min로 공급되었으며 이동상 가스는 초고순도 질소를 사용하였다. Head pressure는 14 psi, split vent는 25 ml/min, septum purge는 2.3 ml/min였다. 190°C의 오븐 온도에서 시료를 주입하여 등온 조건에서 40분간 시료의 지방산 피크를 분석하였다.
알파리놀렌산 합성 유전자 발현제어용 RNAi운반체 제작
들깨 종자의 알파리놀렌산 함량을 낮추기 위해서 알파리놀렌산 생합성의 주된 기능을 하는 소포체 특이적인 FAD3 유전자의 발현을 억제하고자 하였다. 알파리놀렌산 생합성에 관여하는 소포체 특이적인 FAD3와 엽록체 특이적인 FAD7 유전자의 염기서열을 비교하여 종자에서 알파리놀렌산 생합성에 주된 역할을 하는 FAD3 유전자의 발현을 특이적으로 억제할 수 있도록 운반체를 제작하고자 하였다. FAD3와 FAD7 유전자의 염기서열과 아미노산 서열을 비교 분석한 결과 FAD3의 218-281 아미노산 서열에 해당하는 부분의 상동성이 57.8%로 가장 낮았다. 이에 해당하는 812-1005 bp 사이의 193 bp 크기의 전사영역 단편(Fig. 1A)을 RT-PCR로 증폭하고 이를 RNAi 운반체 제작에 사용하였다. PCR로 증폭된 193 bp 유전자 단편을 pDONR201 운반체와 Gateway BP 재조합반응을 거쳐 kanamycin 저항성을 가지는 pENTR-FAD3-193의 중간 클로닝 운반체를 제작하고 클로닝한 유전자 단편의 염기서열을 확인하여 오류가 없는 클론을 선택하여 후속 실험을 진행하였다. 최종 식물형질전환 운반체를 제작하기 위해 발현 억제를 목적으로 하는 유전자 단편을 정방향과 역방향으로 동시에 삽입하면서 두 유전자 사이에 회문(palindrome) 구조를 형성하도록 인트론이 내재되어 있는 pB7GWIWG2(II) 운반체를 선택하였다. 이 운반체는 미생물 선발을 위한 항생제로 spectinomycin 저항성 유전자를 가지며, 식물체 선발마커로 바스타 제초제(Basta™, Bayer CropScience)의 유효성분인 phosphinothricin (PPT)에 저항성을 부여하는 PPT acetyltransferase (PAT or Bar) 유전자를 포함하고 있다. pENTR-FAD3-193 운반체와 pB7GWIWG2(II) 운반체를 Gateway LR 재조합 반응을 거쳐 최종적인 pF3SiA 운반체가 완성되었다(Fig. 1B). pF3SiA 운반체는 BamHI, HindIII, AatII, SacI, KpnI 등의 제한효소 처리를 통해 유전자가 원하는 방향으로 삽입되었음을 확인하였고 염기서열분석을 통해 연결부위의 염기서열을 확인하였다(data not shown). 확인이 끝난 운반체를 식물체에 감염시 발병력(virulence)이 높은 Agrobacterium tumefaciens GV3101 균주에 도입하여 들깨 형질전환에 사용하였다.
Fig. 1
The comparison of target nucleotide sequences between FAD3 and FAD7 of perilla and the construct of RNAi vector to suppress the expression of FAD3 gene encoding Δ15 desaturase. A. Blue arrows represent the DNA region, corresponding to 218-281 aa region where the lowest homology between FAD3 and FAD7. B. pF3SiA vector construct. Asterisk demonstrates unique restriction enzyme site. Bar, phosphinothricin acetyltransferase; CmR, chloramphenicol resistance; F3A, antisense FAD3 fragment for RNAi; F3S, sense FAD3 fragment for RNAi; H, HindIII; K, KpnI; LB, left border; P35S, CaMV 35S promoter; RB, right border; RI, EcoRI; S, SacI; T35S, CaMV 35S terminator.
KJBS_50_463_fig_1.jpg
들깨 형질전환 및 형질전환체 선발
들깨 종자를 살균, 소독 후 petri dish에 20개 정도 파종하여 6-7일 정도된 식물체의 배축을 이용하여 0.5 cm정도 절단 후 pF3SiA가 도입된Agrobacterium을 20분정도 감염시킨 후 3일간 공동배양하고 PPT 1.2 mg/L가 포함된 선발배지에 이식하였다. 치상한 배축은 선발배지에서 2-3주 단위로 새로운 배지에 옮겨주었으며 1-2개월 경과 후 신초가 형성되었다. 형성된 신초를 절단하여 뿌리 유기배지로 이식하여 뿌리를 유도하고 순화하여 형질전환여부를 확인하였다. 이러한 들깨 형질전환과정을 통해서 PPT에 저항성을 가진 재분화식물체 20개체를 확보하여 순화한 후 온실에서 8엽 이상으로 생육이 안정되었을 때 0.3% (v/v) 바스타 제초제를 살포하여 생존한 17개체 들깨 형질전환체를 획득하였고 온실에서 재배하여 T1 종자를 수확하였다(Fig. 2).
Fig. 2
Procedure of perilla transformation using Agrobacterium tumeraciens. A. Cocultivation, B. Shoot induction, C. Shoot elongation, D. Root induction, E. Acclimation, F. Cultivation in Greenhouse.
KJBS_50_463_fig_2.jpg
형질전환을 위해 종자를 살균, 소독시 유지작물의 종자 특성상 종피에 기름기가 많아서 살균이 미흡한 경우가 많아서 Tween-20같은 계면활성제를 이용하여 살균을 하였으나 만족할 만한 결과를 얻지는 못하였다. 종피 살균이 완전히 되지 않는 것 때문에 발아시 세균이나 곰팡이로 오염되는 경우가 상당히 많아 1차 파종 후 건전하게 발아된 개체를 대상으로 2차 배지에 옮겨서 오염을 최대한 방지하면서 형질전환의 재료로 사용하였으나 노력과 시간이 많이 소요되는 것에 비해 효율은 좋지 않았다. 특히 노지에서 재배한 종자를 대상으로 형질전환 실험을 하였을 때는 오염도가 상당히 높았다. 동일한 종자를 사용하여 다양한 살균조건과 배양조건 등을 시도하여 오염을 해결하려고 하였으나 큰 차이는 없었다. 이와는 달리 겨울철 온실에서 재배한 종자의 충실도는 노지에 비해서 낮아서 배축의 굵기가 약하고 발아가 활발하지는 않았지만 기내에서 종자발아시 오염도가 낮아서 형질전환시 높은 효율로 재료를 확보할 수 있는 장점이 있었다. 겨울철 온실 재배하여 수확한 종자의 경우는 노지 재배에 비해 다양한 병원균의 자연 감염이 상대적으로 낮기 때문으로 판단되었으며 유지작물의 형질전환용 종자 확보시 고려할 만한 방법으로 생각되었다.
들깨 형질전환체에서 삽입유전자의 발현 분석
형질전환된 들깨의 유전자 발현분석을 위해 잎과 수정 후 15일된 미숙종자에서 total RNA를 분리하여 Northern blot을 실시하였다. 그 결과 대조구인 엽실들깨 잎에서는 FAD3 유전자의 발현이 되지 않는 것으로 보였으며 종자에서는 FAD3 유전자의 발현량이 높았다. 이 결과는 소포체 특이적인 FAD3 유전자의 발현이 잎에서는 거의 되지 않는 반면 종자에서 상대적으로 높아서 들깨 종자에서 알파리놀렌산 함량이 높은 이유를 보여주었다. 형질전환 여부가 확인된 17개체의 잎과 미숙종자에서 FAD3 발현량을 확인한 결과 잎에서는 대조구와 마찬가지로 전체적으로 발현이 거의 되지 않은 반면 미숙종자에서는 대조구와 유사하게 FAD3의 발현이 강한 개체가 4개체(1, 12, 14, 17), 발현이 중간 정도인 9개체(4, 5, 6, 8, 9, 10, 11, 15, 16) 그리고 대조구에 비해 발현이 많이 약한 4개체(2, 3, 7, 13)를 확인할 수 있었다(Fig. 3). 이는 1, 12, 14, 17 개체를 제외한 나머지 13개체에서는 FAD3 유전자의 발현이 RNAi로 인해 약해졌음을 보여주는 것이다. 유전자 도입 여부를 선발마커인 바스타에 저항성을 보이는 생물반응 여부로만 판단하였기 때문에 유전자의 발현이 강한 4개 개체가 실제로 유전자의 삽입이 되었는데 발현이 제대로 안된 것인지 RNAi 발현 카세트 부분의 삽입에 문제가 있었는지 여부는 Northern blot 실험만으로는 알 수는 없었다. 이를 확인하기 위해서는 추가적인 실험이 필요하였으나 FAD3 유전자 발현이 약해짐으로써 알파리놀렌산 함량이 낮은 들깨를 얻기 위한 목적과는 달라 별도의 확인 과정은 거치지 않았다.
Fig. 3
Northern blot analysis of perilla FAD3 in FAD3-RNAi transgenic perilla. Total RNA was extracted from control plant and transgenic plants as follows : CL, control leaves; CS, control developing seeds; 1-17 L and 1-17S, leaves and developing seeds of 1-17 transgenic line, respectively. The blot was probed with 193 bp FAD 3 sequence by random labelling.
KJBS_50_463_fig_3.jpg
형질전환 들깨종자에서 지방산 조성 분석 및 도입유전자 분리비 검정
유전자의 발현이 확인된 형질전환 들깨 총 17개체 중 고사하였거나 종자량이 적어 지방산 분석이 불가능한 5개체(2, 3, 11, 12, 16번)를 제외한 12개체를 대상으로 종자에서 지방산 조성을 분석하였다. 대조구로 사용한 엽실 들깨의 총 지방산 함량은 422 μg/mg 정도이며 지방산 조성은 팔미트산(palmitic acid, 16:0) 7.5%, 스테아르산(stearic acid, 18:0) 2%, 올레산 13.5%, 리놀레산 16.5%, 알파리놀렌산 60.5%로 구성되어 있다. 형질전환체의 지방산 분석결과 대조구와 비슷한 60%대의 알파리놀렌산 함량을 가진 개체는 3개체(1, 14, 17번)이며 30-40%는 7개체(4, 5, 6, 8, 9, 10, 15번), 20%대는 2개체(7, 13번)였다(Fig. 4). 알파리놀렌산 고함유3개체를 제외하고 알파리놀렌산의 함량이 감소한 것과 달리 리놀레산 함량은 대조구가 16% 인데 비해서 전반적으로 30-40%로 증가하였으며 7번 개체는 57%, 13번 개체는 47%로 상대적으로 높았다. 그 외 팔미트산, 스테아르산, 올레산 함량은 대조구에 비해서 큰 변화가 없었다. 이는 우리가 도입한 FAD3 유전자 단편 RNAi운반체가 FAD3 유전자의 발현을 억제하여 알파리놀렌산 생합성을 저해함으로 전구체인 리놀레산 함량은 상대적으로 증대시키나 다른 지방산 함량은 별다른 영향을 미치지 않음을 알 수 있다(Thambugala et al. 2013). 이와는 달리 콩(Bilyeu et al. 2011), Brassica rapa (Cloutier et al. 2011), 옥수수(Wassom et al. 2008) 등의 fad3돌연변이 식물체에서 올레산의 함량이 증가하는 경우도 보고된 바 있다. 지방산 분석 결과와 유전자 발현분석 결과를 종합한 결과 FAD3 유전자의 발현이 강한 3개체(1, 14, 17번), 발현이 중간 정도인 7개체(4, 5, 6, 8, 9, 10, 15번), 발현이 억제된 3개체(7, 13번)들의 알파리놀렌산 함량이 각각 60%대, 30-40%, 20%대의 개체로 유전자 발현 저해 정도와 알파리놀렌산의 함량은 일관성을 가지고 일치함을 볼 수 있었다(Figs. 3, 4). 알파리놀렌산의 함량을 저하시킴으로 종자유의 산패를 방지하고 고온에서 사용 하기 위해 다양한 시도들이 진행되었다. B. napus의 경우 FAD3 유전자 내에 돌연변이를 이용하여 알파리놀렌산의 함량을 2-3%로 저하시켰으며(Spasibionek 2006) 아마(flax)의 경우는 nonsense- mediated mRNA decay결과로 알파리놀렌산의 함량을 저하시켰다(Vrinten et al. 2005). 형질전환 들깨의 지방산 함량이 대조구에 비해 96-113% 범위에 있어 지방산의 절대량에 큰 변화가 없었으며 표현형에서도 큰 변화는 없었다. 이는 종자특이적으로 발현되는 FAD3의 발현을 억제하였으므로 식물체 전체적인 표현형의 변화가 나타나지 않으며 종자의 저장 지방인 알파리놀렌산 함량만 변화되는 것이라고 생각할 수 있다(Singer et al. 2014). 한편 FAD3 유전자의 전체서열을 역방향(antisense)으로 삽입한 운반체를 이용하여 들깨에 형질전환하여 제초제 저항성을 보이는 6개체를 획득하여 지방산 분석을 수행한 결과 대조구인 엽실 들깨와 비교하여 별다른 변화가 없어서 더 이상의 분석을 수행하지 않았다(data not shown). 역방향으로 유전자의 삽입보다는 RNAi를 이용한 방법이 더 효율적임을 간접적으로 확인할 수 있었다. 형질전환계통의 후대 유전양상을 보기 위해 알파리놀렌산 함량이 가장 크게 감소한 7번과 13번 계통의 T2 종자를 수확하여 지방산 분석한 결과 알파리놀렌산 함량이 7-1계통은 6.0%, 7-2계통은 26.4%, 7-3계통은 20.2%를 나타내었으며 13-1계통은 10.2%, 13-2계통 22.1%, 13-3계통 14.3%를 나타내었다(Table 1). 7번 계통 T1 종자의 알파리놀렌산 함량이 20.4%에서 T2 종자에서는 6.0%, 26.4%, 20.2%로 변화하였으며 특히7-1번 계통은 최대 6%까지 감소함을 보였다. 이 계통의 유전자 도입 분리비를 검정한 결과 최대로 감소한 7-1번 계통은 동형접합체이었으며 7-2와 7-3번 계통은 이형접합체임을 확인할 수 있었다. 마찬가지로 13번 계통도 동형접합체인 13-1번과 13-3번 계통은 10.2%와 14.3%의 알파리놀렌산의 함량을 나타내었다.
Fig. 4
Fatty acid composition of FAD3-RNAi perilla mature seeds. Control is Perilla frutescens cv. Yeobsil plant and 1-17 is transgenic plants. Each bars represents the means ±SD of duplicate independent measurements.
KJBS_50_463_fig_4.jpg
Table 1
Fatty acid compositions of FAD3-RNAi transgenic perilla T2 seeds.
Table 1
Line (T2) The proportions of fatty acid (%) z
Palmitic acid (16:0) Stearic acid (18:0) Oleic acid (18:1) Linoleic acid (18:2) Linolenic acid (18:3) Total FA/seed (μg/mg)
Control (Yeobsil) 8.0±0.8 2.1±0.2 11.9±0.1** 14.9±1.9 63.1±1.2*** 451.8±20.9
7-1 7.6±0.1 1.9±0.1 14.8±1.1** 69.7±4.6** y 6.0±3.3*** 445.4±6.88
7-2 7.3±0.0 1.9±0.0 14.1±0.6* 50.3±1.5** 26.4±0.9*** 482.3±11.4
7-3 7.5±0.0 1.9±0.1 14.3±0.0** 56.1±1.1** 20.2±1.0*** 491.7±10.9
13-1 7.5±0.1 2.0±0.1 14.2±0.3** 66.1±0.5*** 10.2±1.0*** 468.4±25.8
13-2 7.5±0.1 1.9±0.0 14.1±0.9** 54.3±0.2** 22.1±1.0*** 422.3±3.94
13-3 7.6±0.3 2.0±0.1 14.2±0.1** 61.9±2.3** 14.3±1.9*** 447.3±11.4

zThe proportions of fatty acid metyl esters (16:0 to 18:3) are indicated by mole %. Experiments were performed in duplicate, and the mean values ± SD are give.

y*indicates significance at p<0.05,

**indicates significance at p<0.01,

***indicates significance at p<0.001 by Student’s t-test.

7번과 13번 계통의 T2종자를 파종후 바스타 제초제 살포후 분리비를 검정한 결과 7-1, 7-5번은 발아한 53, 55개체가 모두가 생존하여 동형접합계통으로 판정되었으며 7-2, 7-3, 7-4, 7-6번 계통은 3:1의 분리비를 보여 상동염색체 중 한 염색체에만 1 copy가 도입된 이형접합계통임을 확인하였다(Table 2). 마찬가지로 13-1, 13-3, 13-6번 계통은 동형접합계통이며 13-2, 13-4번 계통은 이형접합 계통임을 확인하였다. 이 결과로 세대가 진전되면서 T2종자에서 동형접합계통이 선발되면서 RNA 간섭을 통한 FAD3 유전자 발현이 T1 종자나 T2 종자 중의 이형접합계통보다 강력하게 억제되어 알파리놀렌산 함량이 줄어들었음을 알 수 있었다.
Table 2
The segregation ratios of FAD3-RNAi transgenic perilla T2 plants measured by treatment of Basta herbicide.
Table 2
Line (T2) Resistantance to Basta Susceptibility to Basta Segregation ratio Zygosity
7-1 53 0 homozygote
7-2 54 12 3:1 heterozygote
7-3 50 15 3:1 heterozygote
7-4 57 14 3:1 heterozygote
7-5 55 0 homozygote
7-6 57 14 3:1 heterozygote
13-1 55 0 homozygote
13-2 55 14 3:1 heterozygote
13-3 59 0 homozygote
13-4 57 10 3:1 heterozygote
13-6 56 0 homozygote
들깨는 우리나라에서 재배되어온 대표적인 유지작물로 지질의 함량 중 알파리놀렌산의 함량이 60% 전후로 불포화도가 높아서 산패가 쉽게 일어나는 문제가 있다. 알파리놀렌산은 소포체 유래의 Δ15 desaturase (FAD3)와 엽록체 유래의 Δ15 desaturase (FAD7)에 의해서 합성된다. 엽록체 유래의 FAD7 유전자 발현의 손상없이 종자의 알파리놀렌산 함량을 낮추기 위해 소포체 유래 FAD3 유전자를 RNAi기법을 이용하여 발현을 억제하였다. 재배종인 엽실들깨의 배축을 이용하여 아그로박테리움 매개 형질전환법으로 제초제(바스타) 저항성을 가진 형질전환체 17개체를 획득하였다. 형질전환체는 0.3% (v/v) 바스타 제초제를 이용하여 선발하였으며 Northern blot으로 FAD3 유전자의 발현이 억제되는 것을 확인하였다. 온실에서 수확한 12개체 종자 지방산 함량을 분석한 결과 알파리놀렌산 함량이 10-20% 2개체, 30-40% 7개체, 대조구와 비슷한 60%대 3개체를 획득하였다. 형질전환체의 T2 종자의 분리비와 지방산 조성을 분석한 결과 동형접합체 계통에서 6-10% 알파리놀렌산 함량을 보였으며 이형접합계통은 20-26% 알파리놀렌산 함량을 나타내어 동형으로 고정시 FAD3 유전자 발현이 상당히 강력히 억제됨을 확인할 수 있었다. 들기름의 지방산 중 알파리놀렌산 함량의 감소는 들깨의 산패를 방지하고 감마리놀렌산 등의 고가의 건강기능성 지방산 생산에 활용할 수 있을 것으로 기대된다.
본 연구는 농촌진흥청 국립농업과학원 농업과학기술 연구개발사업(과제번호: PJ01246801)에 의해 이루어진 것임.
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Development of Perilla frutescens with Low Levels of Alpha-Linolenic Acid by Inhibition of a delta 15 desaturase Gene
Korean. J. Breed. Sci.. 2018;50(4):463-471.   Published online December 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.4.463
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Development of Perilla frutescens with Low Levels of Alpha-Linolenic Acid by Inhibition of a delta 15 desaturase Gene
Korean. J. Breed. Sci.. 2018;50(4):463-471.   Published online December 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.4.463
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Development of Perilla frutescens with Low Levels of Alpha-Linolenic Acid by Inhibition of a delta 15 desaturase Gene
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Fig. 1 The comparison of target nucleotide sequences between FAD3 and FAD7 of perilla and the construct of RNAi vector to suppress the expression of FAD3 gene encoding Δ15 desaturase. A. Blue arrows represent the DNA region, corresponding to 218-281 aa region where the lowest homology between FAD3 and FAD7. B. pF3SiA vector construct. Asterisk demonstrates unique restriction enzyme site. Bar, phosphinothricin acetyltransferase; CmR, chloramphenicol resistance; F3A, antisense FAD3 fragment for RNAi; F3S, sense FAD3 fragment for RNAi; H, HindIII; K, KpnI; LB, left border; P35S, CaMV 35S promoter; RB, right border; RI, EcoRI; S, SacI; T35S, CaMV 35S terminator.
Fig. 2 Procedure of perilla transformation using Agrobacterium tumeraciens. A. Cocultivation, B. Shoot induction, C. Shoot elongation, D. Root induction, E. Acclimation, F. Cultivation in Greenhouse.
Fig. 3 Northern blot analysis of perilla FAD3 in FAD3-RNAi transgenic perilla. Total RNA was extracted from control plant and transgenic plants as follows : CL, control leaves; CS, control developing seeds; 1-17 L and 1-17S, leaves and developing seeds of 1-17 transgenic line, respectively. The blot was probed with 193 bp FAD 3 sequence by random labelling.
Fig. 4 Fatty acid composition of FAD3-RNAi perilla mature seeds. Control is Perilla frutescens cv. Yeobsil plant and 1-17 is transgenic plants. Each bars represents the means ±SD of duplicate independent measurements.

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Development of Perilla frutescens with Low Levels of Alpha-Linolenic Acid by Inhibition of a delta 15 desaturase Gene

Fatty acid compositions of FAD3-RNAi transgenic perilla T2 seeds.

Line (T2) The proportions of fatty acid (%) z
Palmitic acid (16:0) Stearic acid (18:0) Oleic acid (18:1) Linoleic acid (18:2) Linolenic acid (18:3) Total FA/seed (μg/mg)
Control (Yeobsil) 8.0±0.8 2.1±0.2 11.9±0.1** 14.9±1.9 63.1±1.2*** 451.8±20.9
7-1 7.6±0.1 1.9±0.1 14.8±1.1** 69.7±4.6** y 6.0±3.3*** 445.4±6.88
7-2 7.3±0.0 1.9±0.0 14.1±0.6* 50.3±1.5** 26.4±0.9*** 482.3±11.4
7-3 7.5±0.0 1.9±0.1 14.3±0.0** 56.1±1.1** 20.2±1.0*** 491.7±10.9
13-1 7.5±0.1 2.0±0.1 14.2±0.3** 66.1±0.5*** 10.2±1.0*** 468.4±25.8
13-2 7.5±0.1 1.9±0.0 14.1±0.9** 54.3±0.2** 22.1±1.0*** 422.3±3.94
13-3 7.6±0.3 2.0±0.1 14.2±0.1** 61.9±2.3** 14.3±1.9*** 447.3±11.4

zThe proportions of fatty acid metyl esters (16:0 to 18:3) are indicated by mole %. Experiments were performed in duplicate, and the mean values ± SD are give.

y*indicates significance at p<0.05,

**indicates significance at p<0.01,

***indicates significance at p<0.001 by Student’s t-test.

The segregation ratios of FAD3-RNAi transgenic perilla T2 plants measured by treatment of Basta herbicide.

Line (T2) Resistantance to Basta Susceptibility to Basta Segregation ratio Zygosity
7-1 53 0 homozygote
7-2 54 12 3:1 heterozygote
7-3 50 15 3:1 heterozygote
7-4 57 14 3:1 heterozygote
7-5 55 0 homozygote
7-6 57 14 3:1 heterozygote
13-1 55 0 homozygote
13-2 55 14 3:1 heterozygote
13-3 59 0 homozygote
13-4 57 10 3:1 heterozygote
13-6 56 0 homozygote
Table 1 Fatty acid compositions of FAD3-RNAi transgenic perilla T2 seeds.

The proportions of fatty acid metyl esters (16:0 to 18:3) are indicated by mole %. Experiments were performed in duplicate, and the mean values ± SD are give.

indicates significance at p<0.05,

indicates significance at p<0.01,

indicates significance at p<0.001 by Student’s t-test.

Table 2 The segregation ratios of FAD3-RNAi transgenic perilla T2 plants measured by treatment of Basta herbicide.

Table 1

Table 2

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