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캘러스 활용도를 향상시키기 위한 벼( L.) 형질전환 시스템 구축

Development of rice(Oryza sativa L.) transformation system to improve callus utilization

Korean Journal of Breeding Science 2017;49(3):170-179.
Published online: August 31, 2017

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*Corresponding Author (hyuns@kribb.re.kr, 042-860-4493, 042-860-4599)
• Received: June 7, 2017   • Accepted: July 31, 2017

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  • Plant molecular farming has attracted a lot of attention lately in the field of mass production of industrially valuable materials by extending application of the plant as a kind of factory concept. Among them, protein expression system using rice(Oryza sativa L.) callus is a technology capable of mass culture and industrialization because of a high expression rate of a target protein. This study was carried out to develop an Agrobacterium-mediated transformation system to increase the utilization of rice callus. The transformation efficiency was improved by using the hand when seeds were de-husked for callus induction. Furthermore, we were possible induction of callus from 6 years old seed smoothly. Selection of the callus contained the target gene was required a cultivation period of at least 3 weeks, and the most efficient selection period was after 6 weeks of culture including one passage. This selection was confirmed that the gene was stably inserted into the genomic DNA of the plant cell by the southern blot analysis and progeny test. Such an efficient selection system of rice callus that can be cultured in the long term will be contribute to the industrialization of useful recombinant proteins using rice.
지난 1990대 초 처음으로 항체와 알부민을 식물에서 발현시킨 이래로(Hiatt et al. 1989, Sijmons et al. 1990) 매우 다양한 식물 종들과 산업용 효소, 성장호르몬, 성장인자들에 이르기까지 폭넓은 대상 단백질로 범위를 확장시켜 나가는 연구들이 줄을 이루었고, 이는 오늘날 분자농업(Molecular farming)이라 칭하 는 특정 연구분야로 자리매김하게 되었다(Obembe et al. 2011, Xu et al. 2012, Joung et al. 2016). 식물을 기반으로 하는 재조합 단백질 생산법은 미생물, 효모, 그리고 형질전환 동물을 이용하는 것에 비해 생산 비용이 적고, 병원성 위해인자들의 오염 위험성이 줄어드는 등 안전한 것이 큰 장점으로 여겨지고 있다(Commandeur et al. 2003, Obembe et al. 2011, Xu et al. 2012). 또한 식물은 소포체, 골지체와 같은 세포 소기관들에 의한 당 단백질의 활성을 위한 기작들이 대장균과 같은 미생물과 비교할 때 동물과 매우 유사하여 당쇄화과정이 필수적인 치료용 단백질의 생산에 적합한 시스템이라 할 수 있다(Ma et al. 2003, Hellwig et al. 2004, Kim et al. 2010). 특히, 식물 세포 내 축적이 아닌 분비되는 단백질의 경우 정제가 간편하다는 장점 또한 있어 더욱 유용하다고 본다(Fischer et al. 2015). 식물세포 의 현탁 배양에 의한 유용단백질의 생산은 온전한 식물체 재배에 비하여 상대적으로 기간이 짧고 제어된 환경에서 지속적인 배양 이 가능하여 경제성과 균일성을 동시에 확보할 수 있다. 이러한 관점에서 볼 때 벼(Oryza sativa L.)는 분자농업에 가장 적합한 식물 중의 하나이다. 더욱이 배지 내 당이 고갈되었을 때 발현되 는 rice alpha amylase 3D (RAmy3D) 프로모터와 세포 밖 분비 신호 서열인 RAmy3D signal peptide 조합을 이용하면 높은 수준의 단백질 발현과 수확이 용이하다(Kim et al. 2008, Shin et al. 2010, Kwon et al. 2013). 벼 종자에서 생산한 알부민, 락토페린, 인슐린(http://www.invitria.com) 등과 벼 현탁배양 세포에서 생산한 트립신, 엔테로키나아제, 성장 인자 (http://www.nbms.co.kr) 등이 이미 산업화되어 연구 시약 및 화장품 성분으로 활용되고 있음은 그 좋은 예이다.
주로 쌍자엽 식물에서 이용되어 오던 Agrobacterium을 이용 한 벼 형질전환 방법은 Chan et al. (1993)이 처음 성공한 이래 Hiei et al. (1994)에 의해 대중적인 방법으로 확립되었으며 그 이후 체세포변이를 줄이고, 보다 단기간에 형질전환체를 만들어 내는 방법 등 많은 연구들이 진행되어 오늘날에는 단자엽 모델식 물로 자리매김하게 되었다(Hiei et al. 1994, Toki 1997, Hiei & Komari 2008, Yang et al. 2013, Clement et al. 2016). 캘러스 유도 초기의 종자(미숙 배)(Toki et al. 2006, Hiei & Komari 2008) 또는 성숙 종자에서 유도된 캘러스(Hiei & Komari 2008, Kim et al. 2009)를 형질전환에 사용하며 항생제 압력에 의한 선발 과정을 거쳐 잎과 뿌리 유도 후 식물체를 얻는 것이 가장 일반적인 과정이다. 이 때, 형질전환에 소모되는 노동력과 시간을 단축하기 위해 Agrobacterium 공동배양과 캘러 스 선발 과정에서 액체 배양을 하거나(Yang et al. 2013), 형질전 환 효율을 향상시키기 위해 Agrobacterium 감염 전 가수분해 효소 전처리와 Agrobacterium 감염 시 열처리를 하는(Clement et al. 2016) 등 형질전환 식물체를 얻기 위한 대부분의 연구들이 진행되어왔다. 앞에서 언급한 벼 캘러스 현탁 배양의 경우, 역시 일반적인 형질전환 방법에 대한 언급만 주를 이루고 있어 분자농 업에의 적용을 위한 캘러스의 고효율 형질전환 시스템은 논문으 로 보고된 바가 없다.
본 실험에서는 기내 배양을 통한 무균 상태의 벼 캘러스 유지와 재조합 단백질의 대량 생산을 위한 벼 캘러스의 효율적인 형질전 환 시스템을 구축하여 벼를 이용한 분자농업 효용성을 제고하고 자 하였다. 이를 위하여 벼 캘러스의 형질전환 후 선발 배지 내에서의 기간을 비교함으로써 최적의 선발 기간을 확립하였다. 그리고 기내 배양을 통한 무균 상태의 벼 캘러스 유지와 배지 내에서 유지한 캘러스로부터 원하는 시기에 언제든지 식물체 재분화를 유도할 수 있는지를 유전자 발현 여부와 후대검정을 통해 확인하였다.
캘러스 유도
본 실험에 사용된 벼(Oryza sativa L. cv. Dong-jin) 종자는 ㈜엔비엠에서 제공받아 사용하였다. 벼 캘러스를 유도하기 위해 서 손 또는 인습기로 종피를 제거한 뒤, 70% 에탄올에 10분간 표면살균을 하고, 3% sodium hypochlorite(Samchun chemical Co., Seoul, Korea) 용액에 10분씩 3회 소독한 후 멸균수에 3회 세척하여 잔여 소독액을 깨끗이 제거하였다. 표면 소독된 종자는 물기를 완전히 건조시켜 배(embryo) 부분이 위쪽을 향하 도록 캘러스 유도배지 N6CI(Table 1)배지에 치상한 후 3~4주간 28°C 암 조건에서 배양하였다(Fig. 1A). 유도된 캘러스는 새로 운 N6CI 배지에 옮겨 3일간 증식시킨 후 형질전환에 사용하였다 (Fig. 1B).
Table 1
Composition of the media used for rice transformation.
Table 1
Step Medium Composition (1L) pH

Callus induction N6CI 3.96g CHU (N6z) medium, 30.0g sucrose, 2.0mg 2,4-D, 0.2mg kinetin, 2.0g phytagel 5.8
Agrobacterium AAMy 2.22g MSx medium, 32.9g sucrose, 18.0g glucose, 0.184g EDTA, 0.25g casamino 5.8
suspension acid, 100uM acetosyringone
Co-cultivation N6CO N6CI medium, 10.0g glucose, 2.0g phytagel, 10.0mg acetosyringone 5.2
Transgenic callus N6SE(.C) N6CI medium, 2.0g phytagel, 50.0mg hygromycin, (500.0mg cefotaxime) 5.8
selection
Callus regeneration N6RE 3.96g N6 medium, 30.0g sucrose, 0.5g glutamine, 0.5g proline, 0.3g casamino acid, 5.8
8g phytagel, 0.5mg NAA, 3.0mg BAP, 250.0mg cefotaxime, 50.0mg hygromycin
Seed germination 1/2 MS 1/2 MS medium, 50mg hygromycin 5.8

zChu et al. (1975)(Cat. No. C0204.0050; Duchefa, Haarlem, The Netherlands).

yHiei et al. (1994).

xMurashige & Skoog(1962)(Cat. No. M0221.0050; Duchefa, Haarlem, The Netherlands).

Fig. 1
Flow chart illustrating the steps for the production of transgenic rice plants. A : Incubation of surface sterilized seeds on N6CI media, B : Callus induction in N6CI media for 3-4 weeks, C : Co-cultivation with Agrobacterium suspension, D : Transformed callus selection on N6SE.C media; E : Histochemical assay of β-glucuronidase in transformed calli, F : In vitro shoot regeneration from transgenic callus on N6RE media, G : Transgenic plant growth, H : Progeny seed germination on 1/2MS medium with 50mg/l hygromycin for 14 days. 3W, 3 weeks; 6W, 6 weeks; 9W, 9 weeks.
KJBS-49-170_F1.gif
Agrobacterium tumefaciens 접종균 준비
벼 형질전환에 사용된 벡터 pCAMBIA1301(CAMBIA, Canberra, Australia)를 열충격(freeze-thaw)방법으로 Agrobacterium tumefaciens LBA4404에 도입하였다. HPT F, 5’-ATG AAA AAG CCT GAA CTC ACC GCG-3’; HPT R, 5’-TCC ATC ACA GTT TGC CAG TGA TAC-3’ primer조합으로 중합효소 연쇄 반응(polymerase chain reaction, PCR)을 수행하여 Agrobacterium 내 벡터 도입을 확인한 후 50mg/l kanamycin, 10mg/l rifampicin, 10mg/l streptomycin이 포함된 YEP고체 배지에 100μl를 도말하였고, 3일간 28°C에서 배양한 뒤 20ml의 AAM(Table 1)액체 배지에 현탁하여(흡광도 650nm=1.8~2.0) 사용하였다.
벼 형질전환 및 형질전환 캘러스 선발
Agrobacterium 현탁액과 준비된 캘러스를 페트리디쉬 (100×20mm)에 같이 넣고 상온에서 60rpm으로 30분간 접종하 였다(Fig. 1C). 접종 후 캘러스는 멸균페이퍼에서 건조하여 잔여 Agrobacterium을 제거한 후 N6CO(Table 1)공동배양배지에 치 상하여 3일간 25℃ 암 조건에서 배양하였다. 공동배양 후 캘러스 는 250mg/l cefotaxime 용액에 3회 세척하고 멸균페이퍼에 올려 물기를 제거하고 50mg/l hygromycin과 500mg/l cefotaxime항 생제를 포함한 선발 배지 N6SE.C(Table 1)로 옮겨 28℃ 암 조건에서 배양하였다. 기본적으로 3주마다 계대배양을 실시하 면서(Fig. 1D), 형질전환된 캘러스의 최적의 선발 기간을 찾기 위하여 배양기간을 3주, 6주(한 번의 계대배양 포함), 및 9주(두 번의 계대배양 포함)로 나누었으며 각 시기마다 효율 비교를 위하여 5-bromo-4-chloro-3-indolyl glucuronide(X-gluc)(LPS solution, Daejeon, Korea)로 염색을 수행하였다(Fig. 1E). 실험 은 3반복으로 진행되었다. 일차배양에서 GUS 염색이 확인된 캘러스는 계대배양에 의한 3주간의 이차배양 후 염색을 수행하 였고, 이 때 동일한 방법으로 염색이 된 캘러스는 다시 계대배양 에 의한 삼차배양 후 3번째 염색을 하여 일차선발시의 결과가 계속적으로 유지되는지 효율 확인을 하였다.
재분화 식물체 선발
형질전환된 캘러스는 재분화 배지N6RE(Table 1)와 25°C, 16시간 광, 광도 125μmol/m2/s의 배양 환경으로 옮겨주어 4주 간격으로 계대배양을 하면서 재분화식물을 유도하였다(Fig. 1F). 각각의 캘러스에서 유도된 재분화 식물체는 한 개체씩 선택하여 50mg/l hygromycin 이 포함된 MS(Table 1)배지에서 생육시켜 분석에 사용하였다. 잎 절편의 X-gluc 염색을 통해 유전자 발현 여부를 확인하였으며, HiYield Genomic DNA Mini kit(RBC Bioscience, Taipei, Taiwan)로 genomic DNA를 추출하여 GUS 유전자 도입 여부를 확인하기 위해 PCR을 수행하였다. PCR에 사용된 primer는 GUS second exon을 특이적으로 증폭 시키는 GUS F, 5’-CTC TAC ACC ACG CCG AAC-3’; GUS R, 5’-GCG CTT GCT GAG TTT CCC-3’ 조합이며 반응 조건 은 95℃에서 2분간 DNA를 변성시키고, 증폭반응(95℃에서 20초, 56℃에서 40초, 72℃에서 45초)을 30회 진행한 후, 최종 신장 반응은 72℃에서 5분간 수행하였다. PCR 증폭 산물은 ecodye(EcoDye DNA staining solution, Biofact, Daejeon, Korea)를 포함한 1% agarose gel상에서 전기영동하여 특이적 밴드를 확인하였다.
형질전환 식물체 분석
배양 기간 6주의 3번 실험군(Table 2) 에서 얻어진 재분화 개체를 이용하여 후대검정을 실시하였다. 기내 배양 식물체를 30℃/20℃(주/야), 16시간 광, 광도 116μmol/m2/s의 생장실에 서 1% Hyponex(Hyponex, Japan)용액으로 일주일간 순화시킨 후 5개월간의 재배 후 T1 종자를 확보하였다(Fig. 1G). 수확한 종자는 물에 깨끗이 씻어서 완전히 말린 후 앞의 방법대로 표면소 독을 실시한 후 50mg/l hygromycin이 포함된 1/2 MS배지에 치상하여 발아를 유도하고(Fig. 1H), 배양 14일 후에 저항성과 감수성 식물체의 분리비를 X²검정에 의해 추정하였다(Seo et al. 2002). 생육중인 식물체 잎 3g에서 cetyltrimethylammonium bromide(CTAB)방법으로 genomic DNA를 추출 한 후(Doyle et al. 1990), 제한효소 HindⅢ(20Unit)(Enzynomics, Daejeon, Korea)를 37℃에서 2일간 처리하였다. 절단된 DNA는 0.8% agarose gel에서 40V로 5시간 전기영동 한 후, Southern (1975) 의 방법으로 positively charged nylon membrane(Roche, Mannheim, Germany)에 DNA를 전이시켰다. 전이된 DNA 단편을 고정하기 위해 UV crosslink(1200×μJ/cm2)를 2회 실시 하였고, PCR digoxigenin(DIG) labeling mix(Roche, Mannheim, Germany)을 이용하여 제작한 GUS probe로 45℃ 에서 16시간 동안 hybridization하였다. Detection과정은 Roche 에서 제공한 chemiluminescent methods로 진행하였고, CDP-Star(Roche, Mannheim, Germany)로 현상하여 결과를 관찰하였다.
Table 2
Selection efficiency of callus by GUS staining that was cultured for 3, 6, or 9 weeks on selection medium with hygromycin and cefotaxime at 28°C in dark.
Table 2
Culture period (weeks) Experiment (replication) GUS staining

1st 2nd 3rd

3 1 0 - -
2 0 - -
3 4 - -

6 1 86.8 100 100
2 100 100 100
3 90.1 98.4 98

9 1 100 100 100
2 91.6 98.99 98.9
3 88.9 100 100
벼 종자로부터의 캘러스 유도
Agrobacterium을 이용한 벼 형질전환에 잎을 이용할 경우 쌍자엽 식물과는 다르게 잎 상처부위의 세포분열이 활발하지 않기 때문에 어려움이 있다(Hiei et al. 2014). 이러한 이유로 대부분 미숙 배나 벼 종자로부터 유도된 캘러스가 재료로 사용되 고 있다(Toki 1997, Lee et al. 2011, Yang et al. 2013, Tran et al. 2015). 본 연구에서도 성숙한 벼 종자로부터 유도된 캘러스 를 형질전환에 사용하였다. 종자를 캘러스 유도 배지에 치상하고 약 3일 후 배 부분에서 싹이 나오기 시작하였고(Fig. 1A), 6일 후 배 부분에서 딱딱한 캘러스가 생성되며, 12일 후 캘러스가 분열하기 시작하였다(Fig. 2C, D). 캘러스의 양과 상태로 보아 형질전환에 사용하기에 적합한 캘러스 유도 기간은 4주가 가장 적당하였다(Fig. 2G, H).
Fig. 2
Efficiency comparison of callus induction caused by seed handling containing de-husking and seed storage duration. Callus induction duration is 2weeks(A, B, C, D) and 4weeks(E, F, G, H). Seed of dong-jin de-husked with hand(A, C, E, G) was more induced callus rather than equipment(B, D, F, H). In case of seed storage duration, seed storaged for 1 year(C, D, G, H) was more vitality than that storaged for 6 years(A, B, E, F). 2W, 2 weeks; 4W, 4 weeks; 6Y, seed storaged for 6 years; 1Y, seed storaged for 1year.
KJBS-49-170_F2.gif
벼 종자로부터 캘러스가 유도되는 비율은 재료로 사용한 종자 의 저장 기간과 종피를 제거하는 방법에 따라 많은 차이가 있었다 (Fig. 2). 6년 보관 종자보다 1년 보관 종자에서 캘러스가 현저히 더 많이 유도되었다. 이전 보고에 따르면 온도와 습도를 조절하지 않은 상태에서 자포니카 벼 종자의 저장 기간에 따른 발아율을 확인했을 때 약 1년 보관 종자는 66.7%가 발아하였고, 약 3년 보관 종자는 거의 발아하지 않았다(Kim et al. 2007). 또한 국내 에서 육성되는 벼 종자를 30개월 저장할 경우 일부 품종에서 평균 40% 이상 활력이 감소하였다(Shon et al. 2014). 벼 종자는 자체적 양분 소모가 있기 때문에 오랜 기간 보관할 경우 종자의 활력이 떨어진다(Kim et al. 2007). 이러한 종자 활력 감소는 캘러스 유도에도 영향을 미치는 것으로 파악된다. 흥미로운 점은 본 연구에서 사용한 6년된 종자도 캘러스 유도율이 양호하다는 점이었다. 이는 종자의 저장 환경이 양호하여 신선도가 유지되었 을 것으로 간주되었다. 또한 종피 제거 시에는 손으로 직접 제거 하는 것이 인습기를 이용할 때보다 캘러스 유도율이 높은 것을 확인할 수 있었다. 인습기를 사용하여 종피를 제거할 때 배 부분 이 반쯤 부러지거나 깎여 나가는 것을 관찰할 수 있었다. 벼 형질전환 시 배의 상태가 건전할수록 형질전환이 성공적으로 이루어진다는 보고가 있다 (Hiei & Komari 2008). 본 실험에서 도 종자의 저장 기간이 짧을수록, 그리고 탈피 시 배 부분에 손상이 가지 않았을 때 캘러스가 많이 유도되었다.
형질전환 캘러스 선발 기간 확립
형질전환 캘러스의 선발 효율을 향상시키기 위하여 Agrobacterium과 공동배양 후 항생제 hygromycin이 포함된 선별배지에서 3주, 6주, 9주 기간 동안 배양하면서 최적의 선발 시기를 GUS단백질 발현을 통해 확인하였다(Table 2). 그 결과, 3주 배양된 캘러스의 경우 염색율 0%의 결과를 보였으며, 한 번의 계대배양을 포함한 6주 배양된 캘러스는 평균 92.3%, positive 결과를 보인 캘러스를 다시 3주 배양한 뒤(총 9주 배양) 확인한 결과 평균 99.5%, 뒤이은 동일 과정 진행 후(총 12주 배양) 확인한 결과 평균 99.3% 염색율을 보여 6주째 보인 결과가 안정적으로 유지됨을 알 수 있었다. 그리고 두 번의 계대배양을 포함한 9주 배양된 캘러스는 9주 째 염색율이 평균 93.5%, positive 결과를 보인 캘러스를 다시 3주 배양한 뒤(총 12주 배양) 확인한 결과 평균 99.7%, 뒤이은 동일 과정 진행 후 평균 99.6%에 달하는 안정적인 결과를 보였다. 종합적으로 보면, Agrobacterium에 의한 형질전환 시 한 번의 계대를 포함한 6주 배양만으로도 원하는 유전자가 도입된 캘러스를 우수한 효율로 선발할 수 있음을 확인하였다. 이는 캘러스로부터 식물체를 재분 화하는 단계를 필요치 않는 캘러스 대량배양에 의해 목표단백질 만을 생산하는 분자농업의 산업화에 실제적으로 적용될 수 있는 간단하지만 효용성 높은 결과라고 판단된다. 다만 다양한 기간 단축 실험이 진행되어 있는 일반적인 벼 형질전환 방법(재분화 식물체를 얻는 경우)에 비해 본 실험에서는 최종 형질전환체(캘 러스)를 얻는 기간이 길다는 단점이 있어 추후 실험에서 형질전 환 캘러스의 선발 기간을 단축시킬 필요가 있다. Yang et al. (2013)은 형질전환 후 선발 배지에서의 캘러스 증식이 2주만에 이루어졌는데, 본 실험과는 달리 캘러스 유도 배지, 캘러스 선발 배지에 L-proline과 myo-inositol이 첨가되어 있었다. L-proline 과 myo-inositol은 캘러스 생육 및 유도율을 증가시키는 역할을 하기 때문에(Lee et al. 2006 , Kwon et al. 1996) 본 실험에서도 캘러스 선발 배지에 L-proline과 myo-inositol을 첨가함으로써 캘러스 유도율을 증가시키면 선발 기간을 6주보다 단축시킬 수 있을 것이라 기대하는 바이다.
위 실험에서 3주 배양된 캘러스의 염색 비율이 0%인 이유는 Agrobacterium과 공동배양 후 형질전환 된 캘러스들에서 새롭게 캘러스가 유도 증식되어 단일군으로 형성되기에는 3주 이상 배양 기간이 필요하기 때문인 것으로 보인다. Toki et al. (2006) 의 연구에서 형질전환 캘러스의 GFP 발현은 선발 배지에서 6~13일 후부터 확인되었다. 하지만 발현 양상이 spot 형태로써 단일 형질전환 캘러스의 증식은 이루어지지 않아 본 실험에서 얻고자 하는 캘러스 단일군을 분리하기에는 어려움이 있어 보인 다. 또한 본 실험에서는 선발 배지에서 3주 배양된 캘러스의 GUS 발현이 0%였는데, 이는 마찬가지로 단일 캘러스의 증식이 덜 이루어진 상태였기 때문에 X-gluc 염색이 진행되었다 하더라 도 형질전환에 사용된 갈변된 캘러스와 X-gluc 염색의 진한 파란색 구분이 어려웠을 것으로 판단된다.
형질전환식물체 재분화 및 분석
캘러스 단계에서의 GUS 단백질 염색에 의한 형질전환 판단 여부를 뒷받침하기 위하여 각 캘러스로부터 재분화 식물체를 확보, 이들의 분자생물학적 분석을 수행하였다. Hygromycin 항생제가 포함된 재분화 배지에 캘러스를 치상하였고, 약 2~4주 후 캘러스의 색이 녹색으로 유도되었다(Fig. 3A). 녹색을 띤 캘러스로부터 벼 지상부가 유도되는 시기는 4~8주로 캘러스 변화 와 유사하게 개체간의 차이가 있었다. Inoue & Maeda(1980)의 보고에 의하면 광조건에서 녹색으로 유도된 캘러스는 잎의 엽록 소 성분과 유사하며, 따라서 이 녹색화 된 캘러스는 엽록체가 발달된 것으로 볼 수 있을 것이다.
Fig. 3
Regeneration of transgenic calli on N6RE medium and expression of GUS gene. A : After calli have cultured for 1 month, color of calli changes yellow to green. Shooting and rooting have induced after calli have cultured for 2 months. Inset shows enlarged change of green calli by arrow, B : PCR detection with genomic DNA of transgenic rice leaves using GUS primers in 1% agarose gel. Product size is 637bp, M, 1kb DNA ladder; X, no template; N, non transgenic callus(Dong-jin); P, plasmid DNA from E.coli, C : GUS staining analysis. The leaves stained with 5-bromo-4-chloro-3-indolyl glucuronide(X-gluc). 3 to 58, independent transgenic lines.
KJBS-49-170_F3.gif
재분화된 식물체의 목적 유전자 도입 여부를 확인하기 위해 50mg/l hygromycin이 포함된 MS 배지에서 생장 후 잎 절편으 로부터 genomic DNA를 추출하여 PCR을 수행하였다. 모든 형질전환체에서 637bp에 해당하는 GUS 특이적 산물을 확인하 였고, 이는 모든 형질전환 식물체의 염색체에 GUS 유전자의 DNA가 삽입되었다는 것을 의미한다(Fig. 3B). 도입된 유전자는 식물체 내에서 전사와 번역을 거쳐 단백질로 발현되는데 이를 확인하기 위하여 X-gluc 염색을 실시하였다 (Fig. 3C). 그 결과 17개체(3, 6, 8, 14, 15, 16, 24, 25, 26, 29, 32, 34, 40, 43, 46, 47, 및 58)에서 GUS 단백질이 뚜렷하게 검출되었다. 일부 식물체에서 나타난 발현 불일치는 몇 가지 이유로 추정될 수 있다. 먼저, 본 실험에서처럼 GUS 염색을 실시하기 전에 개체를 선별한 경우 GUS의 활성은 높으나 염색은 되지 않는 개체가 선별되는 경우가 있다. 그 이유는 일부 세포에 활성을 억제시키거 나 단백질 발현을 억제시키는 GUS 염색 반응에 대한 억제제가 내포되어있기 때문이다(Meijer et al. 1991). 또 다른 이유는 일부 세포에서 GUS유전자 또는 35S promoter와 GUS 유전자 사이에 나타난 메틸레이션에 의해 키메라 발현이 나타났기 때문 이라고 볼 수 있으며 이러한 개체는 향후 형질전환체 선발에서 제외되고 있다(Pavingerova et al. 1997, Ko et al. 1998, Meyer et al. 1992). 또한 GUS 염색에 사용한 잎의 노화로 인한 염색의 어려움이 있다. Li et al. (2017)는 콩의 positive형질전환체를 확인하는 몇 가지 방법 중에서 GUS염색은 뿌리가 유도될 즈음의 어린 식물체 잎에서는 염색이 되나 재배되고 있는 오래된 식물 잎에서는 염색이 되지 않음이 보고된 바 있다.
벼 형질전환체 후대검정
앞에서 확보한 17개 식물체의 도입된 유전자 copy 수 확인을 위해 서던 블롯 분석 결과, negative control(NC)로 사용한 형질 전환 하지 않은 동진벼를 제외한 모든 개체에서 GUS유전자가 1~2 copy로 안정적으로 검출됨을 확인하였다(Fig. 4). 다만, 이들 중 많은 개체들에서 10Kb이상의 위치에서 밴드가 관찰되 었고 이들은 벡터 backbone 서열을 포함하는 위치라고 추측된 다. Agrobacterium을 매개로 하는 형질전환 시 벡터 backbone서 열이 식물체 핵 내로 도입되는 경우는 많이 일어나는 것으로(Ulker et al. 2008) Arabidopsis thaliana 에서 8.3%(Kleinboelting et al, 2015), kumquats는 23.81%(Yang et al. 2017)으로 보고된 바 있다. 명확한 기작은 계속해서 연구중이나 벡터 내의 T-DNA 의 RB, LB부분이 절단되어 T-strand가 식물의 게놈으로 운반되 는데 LB에서 종결을 인식하지 못하고 전사가 계속 진행되어 backbone이 같이 도입되었거나, T-strand가 형성될 때 LB에서 RB 방향으로 진행되어 벡터 전체가 도입되어 발생하는 것으로 알려져 있다(Lim et al. 2008). 또 다른 가능성은 식물체에 여전히 잔재되어 있을 Agrobacterium에 의한 오염이다. Tang et al. (2000)은 호두 체세포배의 75%에서 네 종류의 항생제 추가에도 불구하고 Agrobacterium이 여전히 존재함을 보고한 바 있다.
Fig. 4
Southern blot analysis of 17 transgenic plants and negative control plant(Dong-jin). Each genomic DNA was digested with Hind III and allowed to hybridize with GUS probe. M, 1kb DNA ladder; N, Negative control(Dong-jin); 3-58, Transgenic line.
KJBS-49-170_F4.gif
서던 블롯에 의한 유전자 도입 분석결과 3, 6, 8번과 24, 25, 26은 동일한 패턴을 보였다. 특이하게도 이들은 초기 선발부 터 철저하게 다른 군으로 분류한 각각의 독립 캘러스 유래임에도 불구하고 이러한 결과를 보였다(Fig. 4). 독립적인 형질전환체 3, 8, 24, 32, 및 40번으로부터 T1종자를 확보하여 도입된 유전자 의 안정적인 후대 전이를 확인하였다. 수확한 종자들을 hygromycin이 포함된 1/2 MS배지에 치상 후 발아율을 확인하 였고, 그 결과 32번 개체의 경우 멘델의 독립의 법칙에 따라 정확한 3:1분리 비율을 보였다(Table 3). 이는 단일 copy로 삽입 이 될 경우 멘델의 법칙이 적용되며 안정적인 후대 전이로 이어짐 을 알 수 있으나 보다 정확하게는 T2종자를 확보하여 동일 실험 후 결과를 보아야 알 수 있을 것이다. 반면, 벡터 backbone서열이 포함되어졌을 것으로 예측되는 다른 개체들은 불규칙한 분리 비율을 보였다. 이러한 점들로 미루어 보아 앞에서 언급한 벡터 backbone서열의 검출은 Agrobacterium오염에 의한 것이 아닌 식물 genomic DNA 내 삽입으로 인한 것으로 생각된다.
Table 3
Segregating ratios of resistant F1 progenies to hygromycin.
Table 3
Lines No. of gene copy No. of seed tested No. of planty Separation proportion X 2

HygR HygS

NCz 0 50 0 50 - -
#3 2 90 35 55 7:9 0.86
#8 2 44 16 28 1:1 3.27
#24 2 12 1 11 1:15 0.09
#32 1 100 77 23 3:1 0.21
#40 2 100 5 95 1:15 0.27

Leaves longer than 2cm were classified as resistant. These seedlings grew in 1/2 MS medium containing 50mg/l hygromycin at 24°C under 16h/8h(light/dark) for 12 days. Significant at the 5% level based on X2 tests.

zNegative control.

yHygR, hygromycin resistant; HygS, hygromycin susceptible.

벼 캘러스를 발현 기반으로 하는 유용단백질 생산시스템은 분자농업 중에서도 우수한 것으로 주목받고 있다. 벼의 형질전환 법은 이미 확립되어 어려움 없이 활용되고 있지만 분자농업 측면에서 캘러스를 활용하기 위한 시스템 개선이 필요하였다. 본 실험에서는 종피를 손으로 조심스럽게 제거하여 형질전환 효율을 높여주었다. Agrobacterium 공동배양 후 한차례의 계대 배양을 포함하는 6주간의 배양만으로도 90% 이상 높은 효율의 선발을 할 수 있었다. 종자 20개에서 유도된 캘러스를 이용하여 형질전환을 실시했을 때 새롭게 선발된 개체는 평균 약 79개체였 고, 약 30% 비율로 재분화가 유도되었다. 따라서 본 실험에서 구축한 벼 캘러스 형질전환 시스템을 이용하여 높은 효율로 형질전환 개체를 확보할 수 있고, 재조합 단백질 대량 생산 등 형질전환 캘러스의 활용도를 높일 수 있을 것이라 예상된다.
분자농업은 식물을 일종의 공장개념으로 확대 적용하여 산업 적으로 가치가 높은 유용물질들을 대량생산하는 분야로 최근 많은 각광을 받고 있다. 그 중에서 벼 캘러스를 이용한 단백질 발현 시스템은 대량 배양이 가능하고 목적 단백질의 높은 발현율 로 인한 산업화가 가능한 기술이다. 본 연구는 이러한 벼(Oryza sativa L.) 캘러스의 활용도를 높이기 위한 효율적인 형질전환 시스템을 구축하기 위해 수행되었다. 형질전환에 이용된 종자의 종피 제거 시 손을 이용함으로써 캘러스 유도율을 높여주었고, 6년 정도의 오래된 종자에서도 원활한 캘러스 유도가 가능하였 다. 목적 유전자가 도입된 캘러스의 선발은 최소한 3주 이상의 배양기간을 필요로 하였고 가장 효율적인 것은 한번의 계대를 포함한 6주 배양 후 선발하는 것이었다. 이러한 선발은 유전자가 식물세포의 genomic DNA 안에 안정적으로 삽입되어 이루어졌 음을 서던 블롯 분석 및 후대검정 등을 통하여 확인할 수 있었다. 이러한 장기적이고 대량의 배양이 가능한 벼 캘러스의 효율적인 선발 시스템은 벼를 이용한 유용 재조합 단백질의 산업화에 실속있는 기술로 기여할 수 있을 것이다.
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Development of rice(Oryza sativa L.) transformation system to improve callus utilization
Korean. J. Breed. Sci.. 2017;49(3):170-179.   Published online September 1, 2017
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Development of rice(Oryza sativa L.) transformation system to improve callus utilization
Image Image Image Image
Fig. 1 Flow chart illustrating the steps for the production of transgenic rice plants. A : Incubation of surface sterilized seeds on N6CI media, B : Callus induction in N6CI media for 3-4 weeks, C : Co-cultivation with Agrobacterium suspension, D : Transformed callus selection on N6SE.C media; E : Histochemical assay of β-glucuronidase in transformed calli, F : In vitro shoot regeneration from transgenic callus on N6RE media, G : Transgenic plant growth, H : Progeny seed germination on 1/2MS medium with 50mg/l hygromycin for 14 days. 3W, 3 weeks; 6W, 6 weeks; 9W, 9 weeks.
Fig. 2 Efficiency comparison of callus induction caused by seed handling containing de-husking and seed storage duration. Callus induction duration is 2weeks(A, B, C, D) and 4weeks(E, F, G, H). Seed of dong-jin de-husked with hand(A, C, E, G) was more induced callus rather than equipment(B, D, F, H). In case of seed storage duration, seed storaged for 1 year(C, D, G, H) was more vitality than that storaged for 6 years(A, B, E, F). 2W, 2 weeks; 4W, 4 weeks; 6Y, seed storaged for 6 years; 1Y, seed storaged for 1year.
Fig. 3 Regeneration of transgenic calli on N6RE medium and expression of GUS gene. A : After calli have cultured for 1 month, color of calli changes yellow to green. Shooting and rooting have induced after calli have cultured for 2 months. Inset shows enlarged change of green calli by arrow, B : PCR detection with genomic DNA of transgenic rice leaves using GUS primers in 1% agarose gel. Product size is 637bp, M, 1kb DNA ladder; X, no template; N, non transgenic callus(Dong-jin); P, plasmid DNA from E.coli, C : GUS staining analysis. The leaves stained with 5-bromo-4-chloro-3-indolyl glucuronide(X-gluc). 3 to 58, independent transgenic lines.
Fig. 4 Southern blot analysis of 17 transgenic plants and negative control plant(Dong-jin). Each genomic DNA was digested with Hind III and allowed to hybridize with GUS probe. M, 1kb DNA ladder; N, Negative control(Dong-jin); 3-58, Transgenic line.
Development of rice(Oryza sativa L.) transformation system to improve callus utilization

Composition of the media used for rice transformation.

Step Medium Composition (1L) pH

Callus induction N6CI 3.96g CHU (N6z) medium, 30.0g sucrose, 2.0mg 2,4-D, 0.2mg kinetin, 2.0g phytagel 5.8
Agrobacterium AAMy 2.22g MSx medium, 32.9g sucrose, 18.0g glucose, 0.184g EDTA, 0.25g casamino 5.8
suspension acid, 100uM acetosyringone
Co-cultivation N6CO N6CI medium, 10.0g glucose, 2.0g phytagel, 10.0mg acetosyringone 5.2
Transgenic callus N6SE(.C) N6CI medium, 2.0g phytagel, 50.0mg hygromycin, (500.0mg cefotaxime) 5.8
selection
Callus regeneration N6RE 3.96g N6 medium, 30.0g sucrose, 0.5g glutamine, 0.5g proline, 0.3g casamino acid, 5.8
8g phytagel, 0.5mg NAA, 3.0mg BAP, 250.0mg cefotaxime, 50.0mg hygromycin
Seed germination 1/2 MS 1/2 MS medium, 50mg hygromycin 5.8

zChu et al. (1975)(Cat. No. C0204.0050; Duchefa, Haarlem, The Netherlands).

yHiei et al. (1994).

xMurashige & Skoog(1962)(Cat. No. M0221.0050; Duchefa, Haarlem, The Netherlands).

Selection efficiency of callus by GUS staining that was cultured for 3, 6, or 9 weeks on selection medium with hygromycin and cefotaxime at 28°C in dark.

Culture period (weeks) Experiment (replication) GUS staining

1st 2nd 3rd

3 1 0 - -
2 0 - -
3 4 - -

6 1 86.8 100 100
2 100 100 100
3 90.1 98.4 98

9 1 100 100 100
2 91.6 98.99 98.9
3 88.9 100 100

Segregating ratios of resistant F1 progenies to hygromycin.

Lines No. of gene copy No. of seed tested No. of planty Separation proportion X 2

HygR HygS

NCz 0 50 0 50 - -
#3 2 90 35 55 7:9 0.86
#8 2 44 16 28 1:1 3.27
#24 2 12 1 11 1:15 0.09
#32 1 100 77 23 3:1 0.21
#40 2 100 5 95 1:15 0.27

Leaves longer than 2cm were classified as resistant. These seedlings grew in 1/2 MS medium containing 50mg/l hygromycin at 24°C under 16h/8h(light/dark) for 12 days. Significant at the 5% level based on X2 tests.

zNegative control.

yHygR, hygromycin resistant; HygS, hygromycin susceptible.

Table 1 Composition of the media used for rice transformation.

Chu et al. (1975)(Cat. No. C0204.0050; Duchefa, Haarlem, The Netherlands).

Hiei et al. (1994).

Murashige & Skoog(1962)(Cat. No. M0221.0050; Duchefa, Haarlem, The Netherlands).

Table 2 Selection efficiency of callus by GUS staining that was cultured for 3, 6, or 9 weeks on selection medium with hygromycin and cefotaxime at 28°C in dark.
Table 3 Segregating ratios of resistant F1 progenies to hygromycin.

Leaves longer than 2cm were classified as resistant. These seedlings grew in 1/2 MS medium containing 50mg/l hygromycin at 24°C under 16h/8h(light/dark) for 12 days. Significant at the 5% level based on X2 tests.

Negative control.

HygR, hygromycin resistant; HygS, hygromycin susceptible.