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Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima
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ARTICLE

Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima

Ju-Won Kang1, Dong-Min Kim2, shetty Sang1, Hyun-Sook Lee1, Sang-Nag Ahn1,*

Published online: February 28, 2015

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*Corresponding author (E-mail: ahnsn@cnu.ac.kr, Tel: +82-42-821-5728, Fax: +82-42-822-2631)
• Received: December 16, 2014   • Revised: February 3, 2015   • Accepted: February 3, 2015

© The Korean Society of Breeding Science

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  • We generated 55 introgression lines via backcrosses using Milyang23 as the recurrent parent and Oryza glaberrima as the donor. 141 SSR markers were used to genotype 55 introgression lines. The 55 introgression lines with Milyang23 were evaluated for physiological traits; fresh shoot weight (FSW), fresh root weight (FRW) and dry shoot weight (DSW) under the control and 30% PEG-treated condition. Three lines showing significant difference with Milyang23 were selected for further analysis. Three lines had four, four and two O. glaberrima homozygous segments, respectively. IL9 performed better than Milyang23 in all traits measured in the 30% PEG-treated condition. IL9 possessed four O. glaberrima introgressions on chromosomes 1, 2, 6 and 7. IL12 performed better than Milyang23 in FSW and FRW. IL55 contains two O. glaberrima introgressions on chromosomes 2 and 6. Three lines shared the O. glaberrima segment delimited by markers RM133-RM225 at chromosomes 6. This region corresponds to the QTL region for drought tolerance reported by other previous studies. Although IL9 and IL12 showed improved drought tolerance at the seedling and vegetative stage, they performed poor under the drought stress at the reproductive stage implying that the level of drought tolerance differs according to the growth stage in rice. IL55 had significantly higher no. of the total grain than Milyang 23. This result seems to indicate that IL55 will be a good resource for drought tolerance breeding. The population would be useful in developing drought tolerant lines in the breeding program.
벼는 아시아 지역의 주 식량자원으로 전 세계에서 재배되 는 벼의 90% 이상이 아시아에서 소비되고 있다. 벼는 저온, 염해, 한발 등 비생물적 재해에 의해 피해를 입는데, 특히 한 발은 벼 생산량을 감소시키는 심각한 비생물적 재해 중 하나 다. 아시아에서만 3천4백만 ha의 수도와 8백만 ha의 육도가 빈번한 가뭄 피해를 겪고 있다(Huke & Huke 1997). 따라서, 내한발성 품종의 육종은 관개수의 절약과 가뭄 하에서 안정 적인 수량 유지 및 피해의 최소화라는 측면에서 매우 효율적 인 방법이다.
내한발성 품종의 육성이 어렵고 관련 형질에 대한 유전분석 이 매우 어려운데 이는 한발조건에서 수량관련 형질의 낮은 유전력과 내건성 형질의 효율적인 선발 기준의 부재(Ouk et al. 2006), 내건성 형질의 평가와 검정조건 설정의 문제점을 들 수 있다(Bernier et al. 2007, Courtois et al. 2000, Lafitte et al. 2006, Venuprasad et al. 2007). 그리고 내건성은 양적형 질로 여러 개의 유전자와 환경의 영향을 받기 때문에 그 기작 을 이해하는 것이 어렵기 때문이다. 작물의 내건성 평가는 매 우 어려운데 이는 한발처리 기간 동안 지속적인 수분 공급, 습도와 온도의 조절이 매우 어렵기 때문이다 포장 조건에서 수분 스트레스의 강도는 식물의 근권의 분포에 따라 달라진 다. 포트 실험은 토양수분 포텐셜을 조절하여 스트레스 조건 의 조절이 가능하지만 이 실험은 노동집약적이고 오차가 발 생한다는 점에서 유전분석에 적합하지 않았다. 이러한 면에서 수경재배는 포트 실험보다 우수하다(Kato et al. 2008). 수경 재배 하에서 삼투압 조절을 위해 PEG를 처리하면 수분 스트 레스를 균일하게 조절할 수 있고(Lu & Neumann 1998, Michel & Kaufmann 1973), 식물 뿌리체계의 형태에 상관없 이 뿌리를 낮은 삼투압에 균일하게 노출시킬 수 있는 장점이 있다(Blum 1989, Cabuslay et al. 2002, Michel 1971). Zhang et al. (2006)도 온실조건에서 PEG 처리가 습도와 온 도의 조절이 쉽기 때문에 내건성이 우수한 벼 계통을 선발하 는데 매우 효과적이고 간편한 방법이라고 제시하고 있다.
서아프리카 지역에서 제한적으로 재배되는 아프리카 재배 벼인 O. glaberrima는 병해충저항성, 내건성 등의 생물, 비생 물적 저항성에 관여하는 유용유전자들을 보유하고 있다(www.scienceinafrica.co.za/nerica.htm). 최근 O. glaberrima 의 염 기서열이 밝혀짐에 따라 이들 계통들이 보유하고 있는 다양 한 유전자의 활용이 가능하게 되었다(Wang et al. 2014). O. glaberrima와 O. sativa 간의 교잡후대에서 불임이 관찰되는 데 이 불임현상은 유용 유전자의 이전을 제한하고 있다(Jones et al. 1997). 그럼에도 불구하고 O. glaberrima의 유용한 형 질들이 성공적으로 O. sativa로 이입된 경우가 보고되고 있다 (Jones et al. 1997, Kang et al. 2008, Suh et al. 2005). Bimpong et al. (2011)은 O. glaberrima와 두 개의 인디카 품 종(IR64, IR55423-01) 간의 교잡에서 유래한 여교잡후대 계 통을 한발조건에서 평가하고 한발저항성에 관여하는 다수의 QTL을 탐색하였다. O. glaberrima 유전자는 신규로 탐색된 QTL 유전자의 50-67%를 차지하였는데 주당수량에 관여하 는 두 개의 유전자가 새롭게 탐지되었으며 두 개의 유전자는 2개 지역에서 공통적으로 발견되었다고 보고하고 있다.
벼 분자표지와 유전자지도의 개발로 QTL 분석이 가능하게 되었고 많은 내건성 관련 QTL들이 보고되고 있다(Bernier et al. 2007, Kumar et al. 2007, Lanceras et al. 2004, Swamy et al. 2013, Vikram et al. 2011, Yue et al. 2008). 보고된 QTL들은 뿌리 특성(길이, 두께, 침투 능력 등), 잎 말림과 고 사, 삼투압 조절 등 한발 처리 시에 식물체가 보이는 2차적인 반응과 관련된 특성에 관여하는 QTL이었으며(Lafitte et al. 2006), 한발 조건에서 수량의 안정성에 관여하는 QTL을 보 고한 연구는 거의 없는 실정이다(Suh et al. 2014).
본 연구에서는 밀양23호/O. glaberrima 조합에서 유래된 55개 BC3F8 계통의 유묘기 내건성을 평가하고 내건성이 양 호한 계통을 선발하기 위해 실시하였다.
공시 재료
아프리카 재배벼인 O. glaberrima를 부본으로 통일형인 밀양23호를 모본으로 교잡을 실시하여 F1 개체를 육성하였다. 육성된 F1에 밀양23호를 부본으로 3회 여교배를 실시하여 표 현형적으로 우수한 141개의 BC3F5 계통을 육성하였다 (Kang et al. 2008). 육성된 141계통의 유전자형과 표현형 자 료를 기초로 하여 각 염색체의 부분을 대표하는 단편을 지닌 55계통을 선발하였다. 이 55계통을 이용하여 유묘기와 영양 생장기 내건성을 조사하였다. 양친 중 O. glaberrima는 종자 량이 부족하여 포장조건에서 출수하지 않기 때문에 조사에서 제외하였다.
특성조사
55 이입계통의 유묘기 내건성 관련 5가지 형질을 조사하였 다. 계통 별 건실한 종자를 선발하여 37°C에서 3일간 소독과 침종 후 발아시켰다. 50립의 발아 종자를 가로 30cm, 세로 10cm의 젖은 페이퍼 타월에 치상하고, 밑부분을 제거한 50ml 팔콘 튜브에 넣어 플라스틱 상자에 7일 동안 생육시켰 다.. 발아 10일 후에 건전한 유묘를 선발하고, 홈이 있는 팔콘 튜브 규격의 스펀지를 이용하여 밑부분이 제거된 팔콘 튜브 하나 당 2개의 유묘를 이앙하였다. 뿌리 생장이 저해받지 않 게 하였으며 플라스틱 상자에 팔콘 튜브를 끼워 넣고 1/2 Yoshida 용액(Yoshida et al. 1976)에서 생육시켰다. 상자별 양친과 55계통이 배치되었고 완전임의배치법 6반복으로 실험 하였다. 양액의 pH를 균일하게 유지하기 위해 3일 마다 양액 을 교체하여, pH 5.5~5.8을 유지하도록 하였다. 발아 18일 후에 30% PEG (Polyethylene glycol, 분자량 6000)를 이용 하여 한발 스트레스 처리를 하였다. 유묘기의 생장상 조건은 낮: 13h, 28°C, 밤: 11h, 20°C로 하였다. 안정적인 한발 처리 를 위해 2일 마다 양액을 교체하였으며, pH는 5.7-5.9로 조절 하였다. 한발 처리 5일 후 잎과 뿌리의 생체중과 건물중, 그리 고 엽면적을 조사하였다. 조직의 수분 유실 방지를 위하여 뿌 리와 잎의 생체중 조사는 오전에 실시하였다. 건물중은 55°C 건조기에 2일 이상 건조시킨 후 전자저울을 이용하여 조사하 였다. 처리구와 대조구의 표현형 값을 이용하여 자료를 분석 하였다. 잎과 뿌리의 감소률(Reduction ratio)은 다음의 식을 이용하였다.
R%=CTC×100
R은 감소률(Reduction ratio), C와 T는 각각 대조구 (Control)와 처리구(Treatment) 측정치를 나타낸다. 유묘기 내건성이 우수하다고 판단된 계통에 대해서 상대수분함량 (RWC: Relative Water Content)을 다음의 식을 이용하여 측 정하였다.
RWC%=FWDWTWDW×100
RWC는 상대수분함량, FW와 DW는 각각 생체중(Fresh weight)과 건물중(Dry weight), 그리고 TW는 포화중(Turgid weight)을 나타낸다.
영양생장기에 55계통의 잎말림 정도를 조사하였으며, 1-5 (1: 저항성, 5: 감수성)의 단계를 주었다. 발아시킨 55계통 종 자를 2L 포트에 20립씩 파종하였다. 10일 뒤 건전한 유묘 2 개를 제외한 나머지를 골라내었다. 한발 처리는 파종 후 35일 부터 시작하였으며, 토양수분이 10 cm 깊이에서 5~10%가 될 때까지 처리하였다. 토양 수분이 5~10%가 될 때까지 걸 린 기간은 5~7일 정도였고, 이 기간을 한발 처리기간으로 정 했다. 잎말림 정도를 조사하기 전 2회 처리하였고, 처리마다 토양 수분이 5~10%가 될 때까지 처리 하였으며, 각 처리별 잎말림 정도를 조사하였다(Thanh et al. 2011).
유묘기와 영양생장기 내건성 검정에서 우수한 3계통을 선 발하여 생식생장기 내건성을 평가하였다. 선발된 3계통과 밀 양23호를 비 가림막이 있는 1 m 깊이의 container에 이앙하 였다. 한발 처리는 출수 10일 전부터 출수 후 20일까지 하였 고, 한발 처리 중 잎말림 현상으로 식물체가 고사하는 것을 방지하기 위해 잎말림을 기준으로 두 번의 관수처리를 하였 다. 건조 스트레스 처리 후 지붕의 덮게를 제거한 후 생육조사 를 실시하였다. 임실립과 불임립은 각각 Conutmate MC1000H (Prince)를 이용하여 계측하였다. 선발 계통과 밀양23호의 분 얼수(PN), 수장(PL), 수당립수(SPP), 임실률(FER) 전체립수 (TGN)는 Oh et al. (2009)의 방법을 이용하여 조사하였고 기 타 재배관리도 Oh et al. (2009)와 동일하게 하였다.
DNA 추출, 마커 분석 및 통계분석
55 계통과 양친의 유묘기 잎을 채취하여 Causse et al. (1994)의 방법으로 DNA를 추출하였다. 양친간 다형성을 보 이는 141개의 SSR 마커를 이용하여 55계통의 유전자형을 검 정하였다. PCR 분석과 silver staining은 Panaud et al. (1996) 의 방법에 준해 실시하였다. 연관지도 작성에 사용된 마커의 순서와 마커 간의 거리는 McCouch et al. (2002)의 보고에 준하였다. 자료분석은 SAS 프로그램을 사용하여 밀양23호와 각 계통간 형질의 평균간 차이에 대한 t-검정을 실시하였다.
내건성 형질 변이
유묘기에 밀양23호와 55 계통의 10가지 내건성 관련 형질 을 조사하였다. 뿌리 건물중을 제외한 모든 형질에서 대조구 와 처리구간에 차이를 보였다. 몇몇 계통은 30% PEG 처리구 에서 밀양23호보다 대부분의 형질에서 우수하였다. 이입계통 의 형질별 빈도 분포를 Fig. 1A.에 제시하였다. 밀양23호의 잎 생체중 평균값은 대조구에서 0.362 g 처리구에서 0.128 g 을 보인 반면 이입계통의 잎 생체중은 대조구와 처리구에서 각각 0.282 g - 0.474 g, 0.083 g - 0.196 g의 분포를 보였다. 9개 계통은 처리구에서 밀양23호보다 더 높은 값을 보였는데, 최대 1.6배 이상 높은 값을 보인 계통도 있었다.
Fig. 1.
Frequency distribution. A) Frequency distribution of ten drought tolerance traits in the 55 introgression lines at seedling stage. B) Frequency distribution of leaf rolling scores in the 55 introgression lines at vegetative stage (P1: Milyang23).
KJBS-47-20_F1.gif
밀양23호의 뿌리 생체중 평균값은 대조구와 처리구에서 각 각 0.2570 g, 0.0987 g을 보인 반면, 이입계통은 대조구와 처 리구에서 각각 0.1578 g - 0.3728 g, 0.0807 g - 0.1325 g의 분포를 보였다. 55 이입계통 중 4계통이 밀양23호보다 처리 구에서 더 높은 값을 보였다.
대조구와 처리구에서의 밀양23호의 잎 건물중은 각각 0.118 g과 0.070 g으로 조사되었다. 55 이입계통의 잎 건물 중은 대조구와 처리구에서 0.085 g - 0.158 g과 0.060 g - 0.125 g의 분포를 보였다. 8개 계통이 처리구에서 밀양23호 보다 잎건물중이 무거웠다. 상대수분함량은 4 계통 모두 밀양 23호보다 통계적으로 유의하게 높았다.
영양생장기에 한발 처리를 통해 55 이입계통과 밀양23호 의 잎말림 정도를 조사하였다. 잎말림 정도는 1 - 5의 점수(1: 저항성, 5: 감수성)를 부여하였다. 대부분의 계통은 잎말림 정 도가 밀양23호와 비슷하거나 심하였으나, 몇몇 계통은 밀양 23호보다 양호하였다(Fig. 1B). 밀양 23호의 평균 잎말림 정 도는 3이었고, 이입계통의 잎말림 정도는 1~5의 분포를 보 였다. 처리구에서 밀양23호보다 잎말림 정도가 양호한 계통 은 8개였다.
유묘기와 영양생장기 한발처리에서 내건성 형질이 우수한 계통을 선발하였다. 선발 계통인 IL9, IL12 그리고 IL55는 밀양23호에 비해 잎말림 정도에서 각각 2.1, 1.3 그리고 1로 밀양 23호와 차이를 보였으나 IL41은 밀양23호와 유의한 차 이를 보이지 않았다. 잎말림 정도에서 밀양23호와 차이를 보 인 IL9, IL12 그리고 IL55 3계통의 지상부생체중 감소율을 조사한 결과, 밀양23호는 47.6%를 보인 반면 IL9, IL12, IL55는 각각 33.5%, 27.1% 그리고 35.5%로 밀양23호와 유 의한 차이를 보였다(Table 1).
Table 1.
Mean comparison of traits related to drought tolerance between Milyang23 and selected ILs.
Table 1.
Line Traitsz
 C.R. (%)w
Seedling stage
 Vegetative stagex
FSWT (g)y FRWT (g) DSWT (g) RFSW (%) RFRW (%) RWC (%) LR score RFW (%)
Milyang23 0.12 d 0.09 c 0.07 c 65.3 b 61.7 b 69.2 b 3.5 47.6 ± 3.4 -
IL9 0.19 a 0.13 a 0.10 a 49.0 a 44.9 a 85.1 a 2.1* 33.5 ± 7.7* 93
IL12 0.16 bc 0.12 bc 0.12 bc 44.3 a 54.0 a 81.4 a 1.3*** 27.1 ± 5.1*** 92.3
IL41 0.19 ab 0.15 ab 0.15 b 46.9 a 47.0 a 84.6 a 3.2 - 96.2
IL55 0.16 bc 0.11 bc 0.11 bc 52.6 a 53.5 ab 84.4 a 1*** 35.5 ± 7.4* 97.2

zFSWT: Fresh shoots weight in treated condition, FRWT: Fresh roots weight in treated condition, DSWT: Dry shoots weight in treated condition, RFSW: Reduction rate of fresh shoots weight, RFRW: Reduction ratio of fresh roots weight, RWC: Relative water content, LR score: Leaf rolling score, RFW: Reduction ratio of fresh weight

yNumbers followed by the same letter in a column are not significantly different at the 5% level based on DMRT

xDifference between Milyang23 and each IL

*, ***significant at P≤0.05 and P≤0.001, respectively

wC.R.: Conversion ratio to Milyang23

생식생장기의 내건성 형질 변이
선발된 3계통을 생식생장기에 대조구와 한발 처리 조건에 서 형질을 평가하였다(Table 2). 전체적으로 모든 계통들은 한발조건에서 대조구에 비해 수수에서는 큰 차이를 보이지 않았으나 수당립수, 임실율, 전체영화수가 감소하였다. 한발 조건에서 반복친인 밀양23호와 이입계통을 비교하면, IL12는 출수기가 밀양23호에 비해 약 3주 늦었고 임실률, 수당립수 가 낮았으나 수수와 수장에서는 차이를 보이지 않았다. IL9는 밀양23호에 비해 출수기, 수수와 수장은 차이를 보이지 않았 으나 임실율이 낮았다. IL55는 밀양23호에 비해 수수는 많고 수장은 짧았으나 임실률은 차이를 보이지 않았다. 3계통은 유 묘기와 영양생장기에서 밀양23호에 비해 내건성이 양호하여 선발되었으나 생식생장기 한발조건에서 IL9와 IL12는 밀양 23호에 비해 수당립수, 임실율이 유의적으로 낮았다. 특히 IL12는 임실률이 5%로서 매우 낮았는데 이는 IL12의 늦은 출수기 때문인 것으로 보인다. IL55는 수장, 수당립수 그리고 임실율에서 밀양23호와 유의한 차이가 없는 반면 전체영화수 에서 유의하게 높은 값을 보였는데 이는 수수가 많기 때문인 것으로 보인다.
Table 2.
Comparison of 6 traits between Milyang23 and selected ILs in the control and drought stress at the reproductive stage.
Table 2.
Line Traitz
PN
 PL
 SPP
 FER(%)
 TGN
 DTH
Cy T C T C T C T C T C
Milyang23 11 11 24.5 24.6 181 120 88.7 64.4 1448 793 112
IL9 13 13NS x 25.3 25.9NS 131 95** 65.8 38.3** 862 483** 114
IL12 11 12NS 25.4 25.5NS 125 109NS 59.6 5.6** 771 76** 134**
IL55 12 13* 25.2 23.2 192 115NS 76.4 70.4NS 12111 985 113

zPN: Panicle number, PL: Panicle length, SPP: Spikelets per panicle, FER: Fertility, TGN: Total fertile grain number, DTH: Days to heading

yC: Control condition, T: Treatment condition

xDifference between Milyang23 and each IL

*, **Significant at P≤0.05 and P≤0.01, respectively

NSNot significant

선발 계통의 유전자형
선발된 4계통, IL9, IL12, IL41 그리고 IL55의 유전적 배 경은 반복친인 밀양23호와 유사했다(Fig. 2). 각 계통들은 1, 2, 3, 6, 7, 8번 염색체에 O. glaberrima 염색체단편이 이입되 었고 그 수는 2 - 5개의 분포를 보였다. 4 계통의 밀양23호로 의 회복률은 92.3-97.2%의 분포를 보였다. 선발된 계통 중 IL12는 밀양23호에 비해 출수기가 약 3주 늦었는데 이는 IL12의 염색체 6번 RM587-RM253 부근에 위치한 출수기 관련 QTL인 qdth6의 영향 때문인 것으로 판단된다(Table 2).
Fig. 2.
Graphical genotypes of selected four lines (A: IL9, B: IL12, C: IL41, D: IL55).
KJBS-47-20_F2.gif
본 연구는 육성된 벼 염색체 이입집단의 내건성을 평가하 고 내건성이 증진된 계통을 선발하기 위해 실시되었다. 공시 된 55 BC3F6 계통은 밀양23호/O. glaberrima (IRGC Acc. No. 103544) 조합에서 육성된 141 BC3F5 계통으로부터 선 발되었다. 벼에서 이입집단은 목표 형질의 QTL 분석과 고밀 도지도 작성 등 여러 분야에서 널리 이용되고 있으며 다양한 품종 조합을 이용하여 육성되었다(Ebitani et al. 2005).
AA 게놈의 O. glaberrima는 병해충저항성, 내재해성 등 유용유전자를 보유하고 있으며(www.scienceinafrica.co.za/nerica.htm) 최근 전체 염기서열이 밝혀졌고(Wang et al. 2014) 여러 연구자들이 O. glaberrima의 유용 유전자들이 재배벼로 이전된 계통을 육성하였다(Aluko et al. 2004, Doi et al. 1998, Jones et al. 1997, Kang et al. 2008, Li et al. 2004, Suh et al. 2005).
지금까지 내건성과 관련된 다수의 QTL들이 F2 (Price et al. 1997, Venuprasad et al. 2009a), RILs (Bernier et al. 2007, Venuprasad et al. 2009b, Yue et al. 2008) 그리고 Doubled haploid (Kumar et al. 2007) 집단을 이용하여 보고 되었다. 지금까지 보고된 QTL 중에서 효과가 크고 지속적으 로 탐지된 QTL들은 DTY12.1 (Bernier et al. 2009), DTY1.1 (Vikram et al. 2011), DTY3.1과 DTY2.1 (Venuprasad et al. 2009b) 등을 들 수 있다. Swamy et al. (2011, 2013)는 Adaysel/IR64 조합 후대를 이용하여 4개의 주동 내건성 QTL 을 탐지하였는데, 이들은 대부분 밭 조건에서 수량성을 증진 시키는 QTL들이다. 염색체 1번에는 많은 연구자들에 의해 보 고된 초장, 잎말림 등의 내건성 관련 QTL이 존재한다(Price et al. 2000, Salunkhe et al. 2011, Yue et al. 2008). Price et al. (2000)는 Bala/Azucena 조합에서 육성된 RIL 집단을 이용하여 염색체 1번에 뿌리 생육에 관련된 가뭄 회피 QTL 을 보고하였다. Yue et al. (2008) 은 염색체 1번의 RM472-RM104 지역에 한발 스트레스 조건에서 농업형질에 관여하는 QTL들이 군집되어 있음을 보고하였다. 또한, 한발 조건에서 초장과 분얼에 관한 QTL이 Lanceras et al. (2004) 에 의해 보고되었으며, 잎말림, 잎한발, 수분효율에 관계된 QTL이 Babu et al. (2003)에 의해 보고되었다. 염색체 7번에 도 다수의 내건성 QTL이 보고되었다(Price et al. 2002, Zheng et al. 2000). Bernier et al. (2007)는 염색체 3번에서 생식생장기 한발 스트레스 조건에서 수량 관여 QTL을 탐지 하였고, Yue et al. (2008)는 유사한 지역에서 지엽길이, 지엽 넓이, 잎 한발 그리고 초장에 관련된 QTL들이 군집되어있음 을 보고하였다. 또한, 유사한 지역에서 뿌리 형태에 관여하는 QTL들이 Price et al. (2000, 2002)에 의해 보고되었다.
본 연구에서는 55개 이입계통 중에서 유묘기 내건성이 양 호한 4계통을 선발하였다. 선발된 IL9는 30% PEG 조건에서 밀양23호보다 조사된 모든 형질에서 우수하였다. IL9는 염색 체 1, 2, 6 그리고 7번에 O. glaberrima 단편을 가지고 있었 다. 염색체 1번의 RM472-RM104 지역은 Yue et al. (2008) 이 보고한 한발조건 하에서 농업형질에 관여하는 QTL들이 군집되어 있는 지역과 유사했다. 또한, 이 지역은 한발 조건에 서 초장과 분얼에 관한 QTL이 Lanceras et al. (2004)에 의 해 보고 되었으며, 잎말림, 잎건조, 수분 효율에 관계된 QTL 이 Babu et al. (2003)에 의해 보고되었다. 염색체 7번 RM214- RM11에 이입된 O. glaberrima 염색체 단편의 위치는 다른 연구자(Price et al. 2002, Yadav et al. 1997, Zheng et al. 2000)에 의해 보고된 내건성 QTL 지역과 유사하였다.염색체 2번의 RM106-RM525 근처에 O. glaberrima 단편이 이입된 계통들(IL35, IL37, IL47)은 한발 조건에서 밀양23호와 비교 하여 조사된 형질에서 통계적 유의성이 없었는데(결과 미제 시), 이 결과로 볼 때, IL9의 내건성 증가는 염색체 1번 또는 7번에 존재하는 O. glaberrima 단편의 영향으로 보인다.
내건성에 관여하는 다수의 QTL들이 염색체 3번 단완 말단에 서 탐지되었다. IL12는 염색체 3, 6 그리고 8번에 O. glaberrima 단편을 포함하였는데 염색체 3번의 RM60-RM22 근처에 위 치한 O. glaberrima 이입 단편은 이전 연구자들에 의해 보고 된 QTL 지역과 비슷한 지역이었다(Bernier et al. 2007, Price et al. 2000, 2002, Yue et al. 2008). IL24와 IL40은 염색체 8번에 IL12과 동일한 단편을 보유하고 있었으나, 밀양23호와 통계적인 유의성이 없었다. 이 결과는 IL12의 내건성 증가는 3번 염색체에 위치하는 O. glaberrima 단편 효과로 보여진 다. 아쉬운 점은 55개 계통 중 IL12 외에는 3번 염색체의 RM7-RM282 지역의 단편을 가지는 다른 계통이 존재하지 않아 이 단편의 효과를 평가하는 것이 불가능하였다.
IL41은 염색체 6번과 9번에 총 3개의 O. glaberrima 단편을 포함하고 있었다. 이 지역들 중 염색체 9번의 RM257-OSR28 은 기보고된 내건성 QTL 지역과 비슷했다(Price et al. 2000, Zhang et al. 2006). IL55는 염색체 2번과 6번에 O. glaberrima 의 단편을 보유하고 있다. 이와 같이 계통마다 하나 이상의 O. glaberrima 단편을 보유하고 있기 때문에 어느 염색체단 편이 을 내건성과 관련되어 있는지 파악하는 것이 어려운 상 황이다. 내건성 형질에 관련하는 염색체지역을 분석하기 위해 서는 이들 계통들과 밀양23호 간의 교배를 통한 분리집단의 육성과 평가가 요구된다. 선발된 IL9, IL12, IL41 그리고 IL55의 4계통들은 염색체 6번의 RM133-RM225부근에 공동 적으로 O. glaberrima 단편을 가지고 있었다. 이 지역은 여러 연구자들이 내건성 관련 QTL의 존재를 보고한 지역으로 본 연구에서 선발된 4계통이 모두 이 지역에서 O. glaberrima 단편을 보유하고 있는 것은 매우 주목할 만한 점이다(Bernier et al. 2007, Price et al. 2002).
본 연구에서는 밀양23호의 배경에 O. glaberrima의 특정 염색체단편을 가지는 55 이입계통의 내건성 관련 형질을 조 사하여 변이를 검정하고 내건성이 향상된 4 계통을 선발하였 다. 특히 IL55는 유묘기, 영양생장기 그리고 생식생장기에서 반복친인 밀양23호에 비해 조사된 내건성 형질에서 우수한 특성을 보였으며 내건성 관련 유전자의 분석 및 교배모본으 로 이용될 수 있을 것이다. 이입계통들은 밀양23호의 유전적 배경에 각 계통마다 서로 다른 O. glaberrima 단편이 이입된 계통으로, 이 집단은 O. glaberrima에서 유래된 내재해성 및 작물학적으로 유용한 유전자의 탐색에 효율적인 도구가 될 것이다.
This work was carried out with the support of “Golden Seed Project” (Project No. PJ00993301), Ministry of Agriculture, Food and Rural Affairs (MAFRA), Ministry of Oceans and Fisheries (MOF), Rural Development Administration (RDA) and Korea Forest Service (KFS), Republic of Korea.
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Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima
Korean. J. Breed. Sci.. ;47(1):20-28.   Published online March 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.1.020
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Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima
Korean. J. Breed. Sci.. ;47(1):20-28.   Published online March 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.1.020
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Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima
Image Image
Fig. 1. Frequency distribution. A) Frequency distribution of ten drought tolerance traits in the 55 introgression lines at seedling stage. B) Frequency distribution of leaf rolling scores in the 55 introgression lines at vegetative stage (P1: Milyang23).
Fig. 2. Graphical genotypes of selected four lines (A: IL9, B: IL12, C: IL41, D: IL55).

Fig. 1.

Fig. 2.


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Evaluation of Introgression Lines for Drought Tolerance Derived from a Cross Between the Oryza Sativa Cultivar Milyang 23 and O. glaberrima

Mean comparison of traits related to drought tolerance between Milyang23 and selected ILs.

Line Traitsz
 C.R. (%)w
Seedling stage
 Vegetative stagex
FSWT (g)y FRWT (g) DSWT (g) RFSW (%) RFRW (%) RWC (%) LR score RFW (%)
Milyang23 0.12 d 0.09 c 0.07 c 65.3 b 61.7 b 69.2 b 3.5 47.6 ± 3.4 -
IL9 0.19 a 0.13 a 0.10 a 49.0 a 44.9 a 85.1 a 2.1* 33.5 ± 7.7* 93
IL12 0.16 bc 0.12 bc 0.12 bc 44.3 a 54.0 a 81.4 a 1.3*** 27.1 ± 5.1*** 92.3
IL41 0.19 ab 0.15 ab 0.15 b 46.9 a 47.0 a 84.6 a 3.2 - 96.2
IL55 0.16 bc 0.11 bc 0.11 bc 52.6 a 53.5 ab 84.4 a 1*** 35.5 ± 7.4* 97.2

FSWT: Fresh shoots weight in treated condition, FRWT: Fresh roots weight in treated condition, DSWT: Dry shoots weight in treated condition, RFSW: Reduction rate of fresh shoots weight, RFRW: Reduction ratio of fresh roots weight, RWC: Relative water content, LR score: Leaf rolling score, RFW: Reduction ratio of fresh weight

Numbers followed by the same letter in a column are not significantly different at the 5% level based on DMRT

Difference between Milyang23 and each IL

significant at P≤0.05 and P≤0.001, respectively

C.R.: Conversion ratio to Milyang23

Comparison of 6 traits between Milyang23 and selected ILs in the control and drought stress at the reproductive stage.

Line Traitz
PN
 PL
 SPP
 FER(%)
 TGN
 DTH
Cy T C T C T C T C T C
Milyang23 11 11 24.5 24.6 181 120 88.7 64.4 1448 793 112
IL9 13 13NS x 25.3 25.9NS 131 95** 65.8 38.3** 862 483** 114
IL12 11 12NS 25.4 25.5NS 125 109NS 59.6 5.6** 771 76** 134**
IL55 12 13* 25.2 23.2 192 115NS 76.4 70.4NS 12111 985 113

PN: Panicle number, PL: Panicle length, SPP: Spikelets per panicle, FER: Fertility, TGN: Total fertile grain number, DTH: Days to heading

C: Control condition, T: Treatment condition

Difference between Milyang23 and each IL

Significant at P≤0.05 and P≤0.01, respectively

Not significant

Table 1. Mean comparison of traits related to drought tolerance between Milyang23 and selected ILs.
Table 2. Comparison of 6 traits between Milyang23 and selected ILs in the control and drought stress at the reproductive stage.

Table 1.

Table 2.

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