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배추()에서 분리한 스트레스 관련 후보 유전자 의 특성 분석

Molecular Characterization of a Putatively Stress-related Gene BrTSR53 Isolated from Chinese Cabbage (Brassica rapa)

A-Ram Kim1,†, Hyemin Lim1,†, Sung Han Park2, Hyeonso Ji1, Ung-Han Yoon1, Tae-Ho Kim1, Jung-Il Cho1, Chang-Kug Kim1, Gang-Seob Lee1,*
The Korean Journal of Breeding Science 2014;46(2):143-151.
Published online: May 31, 2014

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*Corresponding author (E-mail: kangslee@korea.kr, Tel: +82-31-299-1656, Fax: +82-31-299-1657)

The first two authors contributed equally to this work.

• Received: December 30, 2013   • Revised: June 9, 2014   • Accepted: June 13, 2014

© The Korean Society of Breeding Science

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  • Crops are exposed to various environmental stresses. These have been affecting the growth of crops, resulting in the severe loss of agronomic production in many countries. Therefore, development of new varieties of resistant crops is required to assure the desired productivity of crops in stress conditions. In this study, a putatively stress-related gene BrTSR53 was isolated from Brassica rapa. The BrTSR53 is 481 bp long and contains ORF region of 234 bp. This ORF showed strong sequence similarities to the uncharacterized genes from Arabidopsis. The expression of BrTSR53 was determined by quantitative real-time PCR analysis. After 3 hr, the highest quantities of mRNA were revealed in cold and salt stress treatments. In drought stress treatments, there was the highest expression after 36 hr. Therefore, it was confirmed that the ORF in BrTSR53 should be a gene that confer increased resistance to B. rapa growing in different stress conditions. The ORF region of BrTSR53 gene was cloned into an expression vector, pYES-DEST52, and a new protein with molecular weight of 13 kDa was detected by western blot analysis. Also, stress tolerance tests showed that BrTSR53-ORF transgenic yeast exhibited increased resistance to the salt stresses compared with the control. In conclusion, the present data predicts that novel ORF in BrTSR53 can serve as an important genetic resource for abiotic stress resistance.
농작물은 기후변화와 환경오염 등에 의한 많은 환경 스트 레스를 받아왔다. 고온, 저온, 가뭄, 고염, 오존, 중금속, 산소 부족 등 다양한 비생물(abiotic) 환경 스트레스는 작물의 성장 과 발달에 영향을 주며, 그 중 가뭄과 고염으로 인한 건조 스 트레스는 전 세계 50% 이상의 경작지의 농업 생산량을 심각 하게 감소시키고 있다(Boyer 1982). 다양한 환경스트레스 중 가뭄, 고염, 저온은 자연환경에서 공통적으로 접해지며 식물이 성장하는데 심각한 영향을 미친다(Xue et al. 2009, Kim et al. 2010).
가뭄 스트레스는 전 세계적으로 작물의 생산량을 감소시키 는 주된 원인이며, 이에 따라 식물의 대응 및 방어 기작에 관한 연구는 그 동안 많은 관심을 받아왔다(Boyer 1982, Cushman & Bohenert 2000). 물 부족 상황에 처했을 때 식물 내에서는 기공개폐, 세포분열 및 생장억제, 광합성 감소 등의 생리학 및 생화학적 반응이 일어난다(Shinozaki & Yamaguchi-sinozaki 2007). 저온 스트레스는 낮은 온도인 0~15°C 사이에서 일어 난다(Lyons 1973). 식물은 성장 단계별 저온에 대한 민감도 가 다르게 나타나는데 일반적으로, 성장과정에서 발아단계와 화분의 성숙 단계가 저온에 민감하다(Patterson et al. 1987, Sataka & Koike 1983). 작물은 고염에 노출되면 세포 내에 삼투압의 불균형(osmotic imbalance)과 이온농도의 불균형 (ionic imbalance)이 일어나게 된다. 또한 2차 스트레스로서 영양장해, 대사장해, 광합성 장해 등이 초래되며, 심할 경우 고사하게 된다(Yun 2005).
하지만 식물은 앞선 환경오염, 지구환경의 급격한 변화로 부터 생존하기 위해 다양한 스트레스에 대한 생존 전략의 일 환으로 체내에 반응 기작을 가지고 있다. 스트레스를 받은 식 물은 많은 유전자들이 발생하고, 유전자 발현조절, 신호전달 의 역할을 한다(Xue et al. 2009). 유전자 발현분석은 생물학 적 연구의 많은 분야에서 중요하며, 생물에서 복잡한 조절 네 트워크의 발생에 유용한 의미를 생산 한다(Chandna et al. 2012). 스트레스에 의해 유도된 유전자 발현은 3개의 그룹으 로 분류된다. 첫 번째로 알고 있는 효소적 또는 구조적 기능 을 가지고 있는 단백질의 유전자, 두 번째는 아직 기능이 알 려지지 않는 단백질 유전자이며 마지막 세 번째는 조절 단백 질이다(Bhatnager-mathur 2008). Chen et al. (2011)의 보고 에 의하면 COR 단백질은 저온 조절 유전자이며, 이 유전자를 이용하여 배추(Brassica rapa)에서는 BrCOR25 유전자를 분 리하였고, 애기장대(Arabidopsis thaliana)에서는 COR15a 및 COR15b 유전자를 분리하였으며, 차나무(Camellia sinensis) 에서는 COR1 유전자를 분리하였다. 또한, Yadav et al. (2012) 의 보고에 의하면, 함초(Salicornia brachiata)에서 유래한 단 백질의 기능이 밝혀지지 않은 SbSI-1 유전자를 분리한 것으 로 보고되었다. 또한 최근에 Jung et al. (2013)의 보고에 의 하면, BrRZFP1 유전자를 저온, 고염 및 건조 스트레스에서 저항성이 증가한 유전자를 배추에서 분리하였으며, Seo et al. (2010)의 보고에서는 BrERF4 유전자를 염과 건조 스트레스 에서 저항성이 증가한 유전자를 배추에서 분리하여 배추유전 자 관련 연구들이 활발히 진행되고 있다.
식물대사의 복잡한 특성은 많은 단백질의 기능 분석을 방해 할 수 있다. 그러나, 효모는 간단한 생물체에서 단백질의 발현은 기능 분석을 위한 효과적인 시스템을 제공한다. Saccharomyces cerevisiae는 단세포로 된 진핵생물이며, 단백질의 정확한 접 힘, 번역 후 변형과 표적 단백질의 유전정보를 통해서 많은 식물 유전자들의 기능 발현이 가능하다(Jellouli et al. 2010). 그러므로 스트레스 저항성 유전자를 효모에 적용하는 것은 산업적으로 중요하다(Hirashi et al. 2006). 보고된 연구에 의 하면, 배추에서 유래한 BrCOR25 유전자와 목화에서 유래한 GhMT3a 유전자를 효모에 형질전환 하여 환경스트레스 저항 성을 확인하였다(Xue et al. 2009, Chen et al. 2011). 본 연 구에서는 배추로부터 저온, 고농도의 염과 가뭄과 같은 환경 스트레스에 관련하여 특이반응을 나타내는 환경스트레스 관 련 후보 유전자인 BrTSR53의 염기서열을 분석하였으며, 환 경저항성에 대한 발현을 확인하기 위한 연구를 수행하였다. 이를 위하여 BrTSR53 유전자의 발현 벡터 시스템을 구축하 였으며, 구축된 벡터 시스템을 Saccharomyces cerevisiae에 형질전환하여 환경스트레스에 대한 발현특성을 분석하였다.
식물재료 및 생육조건
배추 식물체는 지부(Brassica rapa, ‘Chiifu’) 품종을 사용 하였다. 페트리 접시에 3M 종이를 얹고 배추종자를 올려놓았 으며, 25°C, 내부조도 3 lux 배양실에서 3일 간 멸균수에 침 지시켜 배양하였다. 발아한 종자는 원예용 상토로 옮겨 심어 주 었고, 25°C, 습도 93% 조건의 온실에서 3주 동안 배양하였다.
식물체와 효모의 스트레스 처리 조건
배추의 스트레스 처리는 Lee et al. (2008)의 방법에 따라 3주간 성장한 배추를 대상으로 수행하였다. 저온 스트레스는 4°C로 처리한 후 시간대별(0.5, 3, 12, 24, 48시간)로 채취하 였다. 건조 스트레스는 25°C에서 건조 처리한 후 시간대별 (0.5, 3, 12, 24, 48시간)로 샘플을 채취하였고, 고염 스트레스 는 250 mM NaCl 용액을 주입한 후 시간대별(6, 12, 24, 36, 48시간)로 샘플을 채취하였다. 채취한 샘플은 total RNA를 추출한 후 cDNA를 합성하여 분석하였다.
효모의 스트레스 처리는 Jeong et al. (2000)의 방법에 따 라 수행하였다. 형질전환 된 효모를 SC-ura minimal 액체배 지(2% glucose)에 접종하고 30°C에서 2일간 배양 한 후 12,000 rpm에서 5분 동안 원심분리를 하였다. 상층액을 버리 고 SC-ura minimal 액체배지(2% galactose)을 넣고 16시간 동안 배양 한 후, 12,000 rpm에서 5분 동안 원심분리를 하였 다. 상층액을 버리고 10 mM MgSO4 용액에 세척을 하였으 며, 원심분리 하여 배양된 효모를 획득하고, 스트레스 처리를 하였다. 저온 처리는 효모를 YPD 배지에 접종하여 각각 4, 10°C를 5~10일 동안 처리하였다. 염분 처리는 YPD 기본 배지에 NaCl (1.0 1.5, 2.0 M)를 넣어서 수행하였고 건조 처 리는 Micro vac MV-100 (Tony, Japan)을 이용하여 20분간 건조 처리한 후에 멸균수로 적당히 희석 후 YPD 배지에 접 종하여 30°C에서 2일 동안 배양하였다.
Quantitative real time-PCR (qRT-PCR) 분석
배추 식물체의 total RNA은 plant mini kit (Quiagen, USA) 로 추출하였고, Sprint™ RT Complete-oligo (dT) 18 kit (Clontech, USA)를 이용하여 cDNA를 합성하였다. 합성된 cDNA를 SYBR Premix Ex Taq (TaKaRa, Japan) 효소를 사 용하여 95°C에서 30초 초기 반응 후, 95°C에서 5초, 60°C에 서 34초를 40 cycle조건으로 ABI StepOnePlus (Applied Biosystems, USA)를 이용하여 qRT-PCR을 수행하였다. 상 대정량은 Livak & schmittgen (2001)의 2-ΔΔCt 방법으로 측 정하였다. 각 시간대별 BrTSR53 유전자 Ct값에서 β-actin (housekeeping gene) 유전자 Ct값을 빼주어 보정한 후, 각 시 간대별 값에서 대조군(0 hr)의 값을 빼주었다. 그리고 각각의 값을 2의 거듭제곱을 하여 대조군(0 hr)의 값을 1로 두고 나머 지 시간대별에 대한 값을 비교하였다. BrTSR53의 primer는 forward primer; 5’-GGTGACGTCATGGCTTGTGGCT-3’, reverse primer; 5’-ACACGTGAAAAACGAGAGGACAACA-3’ 사용하였고, 비교 유전자 β-actin의 primer는 forward primer; 5’-TGGGCGTACTACTGGTATTGTG-3’, reverse primer; 5’-TGTACCCTCTCTCGGTGAGAAT-3’ 사용하였다.
효모 발현 벡터 구축과 형질전환
BrTSR53 유전자를 효모에서 발현시키기 위하여 pYESDEST52 gateway 벡터 시스템을 이용하였다. BrTSR53 유전 자를 분리하기 위하여 배추 cDNA를 주형으로 RT-PCR을 수행 하였다. TaKaRa Ex Taq™ (TaKaRa, Japan)효소와 BrTSR53의 forward primer; 5’-AAAATGTCTTCATGGAAGAAAACA-3’, reverse primer; 5’-AGAAGACAGATTGTTTGAAGA-3’를 이용하였으며 PCR 반응 조건은 초기(initiation) 95°C에서 5 분 처리 후 95°C에서 30초, 56°C에서 30초, 72°C에서 1분을 총 35회 반복 수행하였으며, 종결(termination) 과정은 72°C에 서 7분간 반응시켰다. pCR8/GW/TOPO 벡터에 sub-cloning을 실시한 클론은 sequence를 확인한 후 pYES-DEST52 Gateway™ 벡터(Invitrogen, USA)에 Gateway LR Clonase™ Ⅱ Enzyme Mix kit (Invitrogen, USA)를 사용하여 효소 발현 벡터에 cloning하였다.
효모에 대한 형질전환은 Saccharomyces cerevisiae (strain W303-1A)를 이용하였으며 YPD (yeast peptone dextrose) 액체배지에 30°C, 200 rpm으로 진탕하면서 2~3일간 배양하 였다. 형질전환은 pYES-DEST52 Gateway™ 벡터의 small-scale 형질전환 방법에 따라 수행하였다. 효모를 YPD 액체배지 5 ml에 접종하여 30°C, 2일간 진탕 배양하였다. 다시 YPD 액 체배지 50 ml 배양액에 3 ml의 cell을 접종하고 30°C, 4시간 진탕 배양하고 4,000 rpm에서 10분간 원심분리를 하였으며, 1X TE 40 ml에 재현탁 시켜준 후 4,000 rpm에서 10분간 원 심분리를 하였다. 상층액을 제거하고 2 ml의 1X LiAC / 0.5X TE에 재현탁 시켜준 후 10분간 상온에 두었다. 재현탁 한 효모 100 μl와 전처리한 salmon sperm DNA 100 μg 및 플라스미드 DNA 1 μg 을 혼합해 주었다. 혼합액에 1X LiAc / 40% PEG-3350 / 1X TE 700 μl를 첨가하여 섞어주고, 3 0°C에서 30분 동안 배양하였다. 88 μl의 DMSO를 첨가하고 42°C에서 7분간 heat shock을 해주었다. 10초 동안 원심분리 후 상층액을 제거하고, 1 ml의 1X TE로 재현탁 시켜주었다. SC-ura minimal 고체배지에 접종하여 도말한 후 30°C, 2일 간 배양하여 콜로니을 확인하였으며, 유전자의 삽입은 PCR 을 통해 확인하였다.
단백질 발현
효모에서 단백질 추출은 Kushnirov (2000)의 방법에 따라 수행하였다. 배양한 효모는 12,000 rpm에서 1분 동안 원심분 리 후 상층액을 버리고, 멸균수를 첨가하였다. 0.2 M NaOH 100 μl를 첨가하여 잘 섞어준 후 상온에서 5분간 두었고, 12,000 rpm으로 1분간 원심분리 하였다. 상층액을 버리고 2X SDS sample buffer를 첨가하였으며, 5분간 끓는 물에서 단백질을 변성 시켜주었다. 이후 SDS-PAGE gel (running gel 15%, stacking gel 1%)을 이용하여 단백질을 분리하였 고, membrane에 transfer 하였다. Transfer된 membrane은 5% skim milk을 첨가하여 blocking하였다. 이후 1차 항체인 anti-V5 (invitrogen, USA)를 1:1000으로 넣어주고 2시간 동 안 반응시켰다. 1X TBS-T (10% Triton X100)를 사용하여 10분간 2회 세척하였으며, 1X TBS를 사용하여 5분간 세척하 였다. 2차 항체인 anti-mouse IgG (Sigma, USA)를 1:10,000 의 비율로 넣어 90분 동안 반응 후 세척과정을 수행하였다. Amersham™ ECL™ Prime Western Blotting Detection Reagent (GE Healthcare, UK)를 이용하여 단백질 발현을 관찰하였다.
스트레스 처리한 배추에서 BrTSR53 유전자의 mRNA 분석
BrTSR53 유전자는 배추의 환경스트레스에 대한 저항성을 microarray 분석으로 찾아낸 저항성을 가진 unknown 유전자 중에 하나이다(Lee et al. 2008). BrTSR53 유전자의 환경스 트레스 저항성을 재확인하기 위해 Lee et al. (2008)와 동일 한 방법으로 배추 식물체를 염분, 저온, 건조 스트레스를 처리 하여 qRT-PCR 분석하였다. 배추에서 염분, 저온, 건조 스트 레스 처리에 따른 BrTSR53 유전자의 mRNA의 발현 결과는 Fig. 1과 같다. 스트레스를 처리하지 않은 대조구를 기준으로 염분 스트레스 처리에서는 30분 처리했을 때는 발현이 감소 하였지만, 3시간 후에는 1.6배까지 증가하였다. 이후 48시간 까지는 점차 감소하는 경향을 보였다. Lee et al. (2013)에 의 하면, 배추 유래 유전자인 BrARP1과 BrRPL27 유전자에 대 하여 염분 처리를 하였을 때, 각각 8시간, 12시간에서 mRNA 양이 가장 높게 증가하였다고 보고하였다. 이 결과와 비교하 였을 때, BrTSR53 유전자는 3시간 염분 처리시에 가장 높은 발현량을 보여 염분 처리에 대하여 상대적으로 빠르게 반응 하는 것으로 나타났다(Fig. 1).
Fig. 1.
Quantitative Real-time PCR analysis of expression of the BrTSR53 gene in Brassica. rapa under salt, cold and drought stress treatment. Rice β-actin was used as a control for relative expression for each treatment. The results are mean values ±SD of three replicates. A. Salt (250 mM NaCl) treatment; B. Cold (4°C) treatment; C. Drought (air dry) treatment.
KJBS-46-143_F1.gif
저온 처리 결과에서는 30분에 2배로 증가하였고, 3시간에 서는 5배의 최대치를 보였으며 12시간 이후부터는 감소하는 경향을 보였다(Fig. 1). 이러한 경향은 BrTSR53 유전자의 염 분 처리한 결과와 유사한 경향을 보였다. Wang et al. (2012) 은, 4°C 저온처리를 하였을 때, 배추(Brassica campestris)에 유래한 BcWRKY46 유전자는 2시간 까지 mRNA 양이 증가 하였고, BrCBF 유전자는 4시간까지 증가하는 것을 보고한 바 있다. 스트레스 관련 후보유전자인 BrTSR53 유전자의 반 응도 3시간에서 최대치를 보이고 있어 유사한 경향을 보였는 데, 이는 분리된 유전자의 유래가 배추임에 따른 결과로 보여 진다.
건조 처리 결과에서는 염분 처리, 저온 처리와 다르게 6시 간에서는 1.9배 증가하였고, 24시간 처리한 결과에서는 0.5배 까지 감소하였으나, 36시간 처리 후 다시 2배로 증가하여 최 대치를 보이는 경향이 나타났다(Fig. 1). Yadav et al. (2012) 의 보고에 의하면, 함초(Salicornia brachiata)에 유래한 SbSI-1 유전자는 건조 처리를 하였을 때, 24시간 까지 mRNA 양이 증가하였다. 이 유전자와 비교하여 보았을 때, BrTSR53 유전 자는 36시간에서 mRNA 양이 증가한 것으로 나타남으로써, 건조에 대해서는 다른 스트레스에서 보다는 상대적으로 늦게 반응이 나타나는 것을 확인되었다.
이 결과들을 종합할 때, 배추에서 유래한 BrTSR53 유전자 는 저온과 염분 등의 스트레스 초기에 발현하는 유전자임을 추론할 수 있었으며, 염분 또는 건조 스트레스 처리보다 저온 처리에서 유전자 발현량이 증가하는 특성을 확인할 수 있었다. 이는 Dai et al. (2007)의 보고에서도 벼에서 유래한 OsISAP1 유전자가 염분, 건조, 저온 스트레스 중 저온 스트레스에서 mRNA 양이 가장 증가한 특성을 보인 것으로 보고되어 비슷 한 경향을 보이는 결과를 얻었다.
BrTSR53 유전자의 염기서열 분석
배추 식물체에서 염분, 저온, 건조 스트레스에 저항성을 가지는 것으로 확인된 신규유전자를 BrTSR53 (Triple Stress Resistance 53 gene)이라고 명명하였다. BrTSR53 (gene number: BRAS0-001S00024153) 유전자의 염기서열 분석결과 mRNA는 481 bp로 구성되며 ORF는 234 bp 되어있다(Fig. 2). 또한 BrTSR53 77개의 아미노산으로 이루어진 단백질을 지령할 수 있다(Fig. 2). 분석된 염기서열을 대상으로 blast 분석결과, Eutrema salsugineum (XM006393369.1)와는 max score가 총 392가 일치하였으며 97%의 상동성을 보여 BrTSR53와 가장 유사한 유전자로 확 인되었다. 그 다음 Arabidopsis lyrata (XM002891379.1)와는 max score가 총 356이 일치하였으며 94%의 상동성을 보였고, 이 외에 Arabidopsis thaliana (CP002684.1), (NM103730.4), (BT011259.1), (BT010781.1), (BX818303.1)와 (AC007932.3) 는 max score는 329이었으며, 91% 상동성을 보이는 것으로 나타났다. 검색결과에서 BrTSR53 유전자는 NCBI에 등록되 어 있지도 않았으며, 이에 따른 주요기능 또한 보고된 바 없 음을 확인할 수 있었다. 그러나 Yang et al. (2013)의 보고에 의하면, ORF에 대한 상동성 분석에서 가장 유사한 Eutrema salsugineum는 A. thaliana 와 Brassica 속과 분류학적으로 거리가 가까운 염생 식물이며, 염분스트레스의 저항성을 연구 하기 위한 모델 식물체인 것으로 알려져 있다. 따라서 BrTSR53 유전자는 그 기능은 밝혀져 있지는 않지만, 염분 스트레스와 관련된 유전자 임을 추정할 수 있었다(Fig. 3).
Fig. 2.
Sequence analysis of the BrTSR53 gene. Full-length nucleotide sequence and amino-acid sequences for ORF region of the BrTSR53 gene isolated from Brassica rapa.
KJBS-46-143_F2.gif
Fig. 3.
Phylogenetic analysis for the BrTSR53 gene isolated from Brassica rapa. The distance was calculated by Kimura’s twoparameter methods and the tree was constructed by using the neighbor-joining method. Number indicates bootstrap values after 1000 replications.
KJBS-46-143_F3.gif
효모에서 BrTSR53 유전자의 발현
BrTSR53 유전자의 기능을 알아보기 위해 효모 단백질 발 현 벡터인 pYES-DEST52 벡터에 BrTSR53 유전자의 ORF를 cloning하였다. BrTSR53 유전자의 삽입위치는 T7 promoter 와 V5 epitope 사이에 위치 하였으며, 단백질 검출을 위한 항 체인 V5 epitope, 단백질 정제를 위한 6X hig tag과 선별마커 인 URA3 합성 유전자로 구성되었다(Fig. 4). pYES-DEST52/BrTSR53 벡터를 Saccharomyces cerevisiae에 형질전환을 수행하였고, PCR 방법으로 형질전환 유무를 확인하였다. 형질 전환체 효모의 단백질을 발현을 확인하기 위하여 SDS-PAGE 와 western blot을 수행하였다. pYES-DEST52 벡터만 삽입한 효모로부터 단백질 발현을 유도(induction)한 경우와 유도하 지 않는 경우(non-induction), pYES-DEST52/BrTSR53 벡터 시스템을 삽입한 효모에 대하여 단백질을 유도한 경우와 유 도하지 않은 경우 등 특정 단백질 발현을 확인하기 위한 4가 지의 조건을 비교하였다. Western blot 분석법을 이용하여 BrTSR53 단백질이 10~15 KDa 사이에서 형성되었음을 확 인할 수 있었다(Fig. 4). 이로써, 효모를 이용한 발현 시스템 에서 BrTSR53 단백질이 발현됨을 알 수 있었으며, 형성된 단백질은 오직 BrTSR53 유전자에 의해서만 형성될 수 있는 특정 단백질임을 확인할 수 있었다.
Fig. 4.
Generation of the BrTSR53 transgenic yeast. A. pYES-DEST54/BrTSR53 vector construct for expression of the BrTSR53 in yeast B. Western blot analysis of the BrTSR53 for protein expression in Saccharomyces cerevisiae; (a) non induction of empty vector; (b) induction of empty vector; (c) non induction of pYES-DEST54/BrTSR53; (d) induction of pYES-DEST54/BrTSR53.
KJBS-46-143_F4.gif
환경스트레스에 대한 저항성
BrTSR53 유전자의 저항성을 알아보기 위해 효모발현시스 템을 이용하여 환경 스트레스를 처리하였다. 대조구인 pYESDEST52 벡터만 삽입한 효모와 pYES-DEST52/BrTSR53 벡 터를 삽입한 효모의 발현을 유도하였으며, 저온, 건조, 염분 스트레스 처리를 하였다.
저온과 건조 처리에서 pYES-DEST52 효모와 pYES-DEST52 /BrTSR53 삽입한 효모는 차이가 보이지 않았다(data not shown). 염분 처리 결과에서는 1 M NaCl 처리에서는 pYES-DEST52 효모와 pYES-DEST52/BrTSR53 효모에서는 차이가 나타나 지 않았으나 1.5 M NaCl 처리에서는 pYES-DEST52 효모보 다 pYES-DEST52/BrTSR53 효모에서 집락이 증가하였다. 또한 2 M NaCl 처리에서는 pYES-DEST52 효모에서는 집 락이 형성되지 않았지만, pYES-DEST52/BrTSR53 효모에서 는 10-2까지 배양된 것을 확인할 수 있었다(Fig. 5). Diao et al. (2010)의 보고에 의하면, Limonium bicolor로부터 유래한 환경스트레스 저항성 유전자인 LbGST1의 형질전환체 효모를 NaHCO3, 건조, 저온, 염분, 고온 및 UV (Ultraviolet radiation) 의 다양한 스트레스 처리를 하였고, LbGST1 유전자를 삽입 한 형질전환체 효모와 유전자를 삽입하지 않은 효모의 세포 를 희석하여 비교하였다. 그 결과 대조군을 제외한 NaHCO3, 건조, 저온, 염분, 고온 및 UV 스트레스를 처리하였을 때, LbGST1의 유전자를 삽입한 형질전환체 효모에서 저항성을 나타내었다. 이 결과와 비교하였을 때, BrTSR53유전자는 효 모발현시스템에서 염분 스트레스에 대해 저항성을 보이는 것 으로 나타났다.
Fig. 5.
Salt stress treatment of the BrTSR53 transgenic yeast cell. pYES-DEST52 and pYES-DEST52/BrTSR53 transgenic yeast cells were spotted on plates with normal growth medium (YPD) or medium containing 1.0 ,1.5 and 2.0 M NaCl, respectively. Transgenic yeast cells were diluted by 10~10-4. The inoculated transgenic yeast cells were incubated at 30°C for 3~5 days.
KJBS-46-143_F5.gif
농작물은 다양한 외부 환경스트레스에 노출되어 있다. 환 경스트레스는 작물의 성장에 영향을 주어 세계 각 지역의 농 업 생산량을 심각하게 감소시키고 있다. 따라서 작물의 생산 성을 높이기 위해서 다양한 환경스트레스에 내성이 강한 새 로운 품종의 개발이 요구된다. 최근의 연구 동향은 환경스트 레스 저항성 유전자를 작물에 도입시켜 환경 변화에 대한 저 항성이 강한 작물을 개발하는 것이다.
본 연구에서는 배추의 저온, 고농도의 염과 건조 등의 환경 스트레스에 대한 저항성 유전자로 추정되는 BrTSR53의 염기 서열을 분석하였다. BrTSR53의 유전자의 총 길이는 481 bp 이며 이중에서 ORF 부위는 234 bp이었다. 이 ORF의 염기서 열 상동성을 분석한 결과 Arabidopsis에서 보고된 유전자와 유사한 것으로 나타났다. BrTSR53의 발현을 분석하기 위하 여 quantitative real-time PCR을 실시하였다. 그 결과 배추를 고염 처리, 저온 처리하고 3시간 후에 가장 높은 mRNA 양을 보였으며, 건조 처리에서는 36시간 후에 발현량이 최대치를 보였다. 따라서 이 ORF는 환경스트레스에 대한 배추의 저항 성 유전자임을 확인하였다.
그리고 BrTSR53 유전자를 효모발현 벡터인 pYES-DEST52 에 삽입하고 western blot 분석법을 통해 효모에서 분자량이 약 13 kDa인 저항성 단백질의 발현을 확인하였다. 또한 BrTSR53 형질전환 효모는 염분 스트레스에 대한 저항성이 증가한 것 으로 나타났다. 따라서 BrTSR53 유전자는 농작물의 환경스 트레스 저항성을 높여줄 수 있는 주요한 유전자원으로 이용 될 수 있다고 사료된다.
본 연구는 농촌진흥청 어젠다(과제번호: PJ010086)와 차세 대 바이오그린 21사업(과제번호: PJ0080912012)의 지원에 의해 이루어진 것임.
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Molecular Characterization of a Putatively Stress-related Gene BrTSR53 Isolated from Chinese Cabbage (Brassica rapa)
Korean. J. Breed. Sci.. 2014;46(2):143-151.   Published online June 30, 2014
DOI: https://doi.org/10.9787/KJBS.2014.46.2.143
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Molecular Characterization of a Putatively Stress-related Gene BrTSR53 Isolated from Chinese Cabbage (Brassica rapa)
Korean. J. Breed. Sci.. 2014;46(2):143-151.   Published online June 30, 2014
DOI: https://doi.org/10.9787/KJBS.2014.46.2.143
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Molecular Characterization of a Putatively Stress-related Gene BrTSR53 Isolated from Chinese Cabbage (Brassica rapa)
Image Image Image Image Image
Fig. 1. Quantitative Real-time PCR analysis of expression of the BrTSR53 gene in Brassica. rapa under salt, cold and drought stress treatment. Rice β-actin was used as a control for relative expression for each treatment. The results are mean values ±SD of three replicates. A. Salt (250 mM NaCl) treatment; B. Cold (4°C) treatment; C. Drought (air dry) treatment.
Fig. 2. Sequence analysis of the BrTSR53 gene. Full-length nucleotide sequence and amino-acid sequences for ORF region of the BrTSR53 gene isolated from Brassica rapa.
Fig. 3. Phylogenetic analysis for the BrTSR53 gene isolated from Brassica rapa. The distance was calculated by Kimura’s twoparameter methods and the tree was constructed by using the neighbor-joining method. Number indicates bootstrap values after 1000 replications.
Fig. 4. Generation of the BrTSR53 transgenic yeast. A. pYES-DEST54/BrTSR53 vector construct for expression of the BrTSR53 in yeast B. Western blot analysis of the BrTSR53 for protein expression in Saccharomyces cerevisiae; (a) non induction of empty vector; (b) induction of empty vector; (c) non induction of pYES-DEST54/BrTSR53; (d) induction of pYES-DEST54/BrTSR53.
Fig. 5. Salt stress treatment of the BrTSR53 transgenic yeast cell. pYES-DEST52 and pYES-DEST52/BrTSR53 transgenic yeast cells were spotted on plates with normal growth medium (YPD) or medium containing 1.0 ,1.5 and 2.0 M NaCl, respectively. Transgenic yeast cells were diluted by 10~10-4. The inoculated transgenic yeast cells were incubated at 30°C for 3~5 days.

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Molecular Characterization of a Putatively Stress-related Gene BrTSR53 Isolated from Chinese Cabbage (Brassica rapa)

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