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통일형 벼 이앙 후 유묘기 재배안정성 증진을 위한 저온생장성 관련 QTL 탐색

장성규1, 이지윤2, 강주원1, 권영호1, 이소명1, 이샛별1, 조준현1, 박동수1, 이종희1, 권순욱3, 조수민1,*

Identification of QTLs Related to Plant Growth at Low Temperatures in the Seedling Stage of Tongil Type Rice after Transplanting

Seong-Gyu Jang1, Ji-Yoon Lee2, Ju-Won Kang1, Youngho Kwon1, So-Myeong Lee1, Sais-Beul Lee1, Jun-Hyeon Cho1, Dong-Soo Park1, Jong-Hee Lee1, Soon-Wook Kwon3, Sumin Jo1,*
Korean Journal of Breeding Science 2024;56(3):225-235.
Published online: September 1, 2024

1농촌진흥청 국립식량과학원 남부작물부 논이용작물과

2농촌진흥청 국립식량과학원 기획조정과

3부산대학교 식물생명과학과

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*Corresponding to Sumin Jo TEL. +82-55-350-1164 E-mail. tnals88319@korea.kr
• Received: May 11, 2024   • Revised: July 25, 2024   • Accepted: July 31, 2024

Copyright © 2024 by the Korean Society of Breeding Science

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  • In Asia, where climate change is increasing the damage caused by cold stress, it is crucial to cultivate varieties with enhanced cold tolerance. In this study, the Tongil variety ‘Hanareum2’ was crossed with the Japonica variety ‘Unkwang’ to improve plant growth ability at low temperatures during the seedling stage. This led to the development of 234 recombinant inbred line (RIL) populations, and a linkage map was constructed using 249 single nucleotide polymorphism (SNP) markers. The RIL populations were transplanted to the field one month earlier than the standard transplanting period, and plant height (PH), leaf number (LN), and dry weight (DW) were measured to identify quantitative trait loci (QTL) associated with plant growth ability at low temperatures during the seedling stage. QTLs related to cold tolerance, particularly those carrying the ‘Unkwang’ allele, were identified in the PH and DW traits. For PH, the QTLs qPH1, qPH5, and qPH8 were located on chromosomes 1, 5, and 8, respectively. Regarding DW, the QTLs qDW1, qDW8, and qDW9 were identified on chromosomes 1, 8, and 9. For the LN trait, qLN3 carrying the ‘Hanareum2’ allele was located on chromosome 3. If the identified QTLs are utilized, they can be incorporated into breeding programs for plant growth at low temperatures during the seedling stage.
  • Keywords: growth ability; low temperature; quantitative trait loci; tongil type; rice
쌀(Oryza sativa L.)은 전 세계 인구 절반이 소비하고 있는 작물이며, 열대 및 아열대 지역이 원산지이기 때문에 온도에 민감한 특성을 가지고 있다. 특히, 저온에서는 생산량이 크게 감소하는 것으로 보고되고 있으며, 전 세계적으로 저온 피해 재배지역은 약 1,500만 ha로 보고되었다. 벼가 재배되는 25개국에서는 저온 피해가 주요 기후 문제 중 하나로 대두되고 있다(Bai et al. 2016, Cruz et al. 2013, Thapa et al. 2020).
4월에 발생하는 한류로 인해 초기 벼 모종이 썩어 종자가 많이 손실되고 성장 기간이 지연되는 경우가 많이 발생한다. 1980년과 1993년에 대한민국에서도 냉해가 발생하여 각각 전년도 대비 국내 벼 재배면적의 약 26%, 9.2%의 수확량 감소를 가져왔다(Lee 2001).
벼 재배에서 최적 온도는 25~30℃이며, 벼의 생육 초기에 15℃ 이하로 온도가 낮아졌을 때 저온스트레스가 발생하며, 종자의 발아가 늦어지고, 유묘기 적고 현상이 발생하는 등 생육에 부정적인 영향을 끼치는 것으로 보고되었다(Yoshida 1981, Zhang et al. 2014). 수잉기와 출수기, 등숙기에도 17~20℃ 이하로 떨어질 시 이삭의 길이가 짧아지고, 종자의 등숙률이 떨어져 수확량이 감소하는 것으로 보고되고 있다. 또한 이는 종자의 발아 불량으로 못자리 육묘 실패, 직파재배 시 입모 부족 등 악영향을 주는 것으로 확인되었다(Hyun et al. 2016, Liu et al. 2018, Ma et al. 2015, Sales et al. 2017, Shim et al. 2022).
Deng et al. (2018)은 Gr89-1×Shuhui527의 재조합자식계통(Recombinant Inbred Lines, RIL)을 활용하여 저온스트레스에 관한 QTL분석을 진행하였다. ‘Gr89-1’은 다년생 벼로서 월동 후에서 그루터기에서 새로운 잎이 자라나 수확할 수 있는 품종으로 내냉성이 강한 품종이다. 해당 형질에 연관된 QTL을 탐색하기 위해 5~22℃의 조건 속에서 286개의 RIL 계통과 198개의 SSR (Simple Sequence Repeat)마커를 통해 3개의 QTL (qOSR2, qOSR3, qOSR8)을 탐색하였다. Mao et al. (2015)은 야생벼 ‘Dongxiang’의 내냉성 관련 QTL을 탐색하고자 RIL집단을 육성하였으며, 총 13개의 QTL을 탐색하였다. 이들 중 3, 7, 8, 11번 염색체에서 주동 QTL이 탐색되었다. Sun et al. (2018)은 저온생장성이 강한 품종인 ‘Kongyu131’을 감수성인 ‘Dongnong422’와 교배하여 190개의 RIL집단을 제작하였고, Bulked Segregant Analysis (BSA)를 통해 저온 스트레스에 관한 3개의 QTL (qPSST5, qPSST6, qPSST7)을 탐색하였다.
본 연구는 저온생장성과 관련된 QTL을 탐색하고자 내냉성이 약한 통일형 벼 ‘한아름2호’와 내냉성이 강한 자포니카 벼 ‘운광’을 이용하여 육성된 234개의 RIL집단을 활용하였다. 우리나라는 온대지역으로 유묘기 단계에서 저온스트레스 내성은 통일형 초다수성 품종의 안정적 생산을 위해 매우 중요한 특성이다. 유묘기 저온생장성에 관련 양적 형질의 탐색과 이를 활용한 통일형 초다수 품종의 내냉성 개량은 향후 기후변화 대응 식량안보 구축에 도움이 될 것으로 기대된다.
한아름2호×운광 RIL 집단 육성 및 특성 조사
본 연구에서 통일형 초다수성 벼의 조기 이앙에 따른 저온생장성에 관련된 QTL을 탐색하기 위하여, 통일형 ‘한아름2호’를 모본으로 하고, 저온스트레스에 강한 자포니카 고품질 품종 ‘운광’을 부본으로 사용하여 2009년 인공교배를 실시하였다. 2010년 F1 식물체를 양성하였으며, F2세대부터 1개체 1계통 육종법(Single Seed Descent, SSD)을 통해 F6 세대까지 세대진전하여 총 234계통의 RIL을 육성하였다. 저온생장성 조사를 위해 모부본과 RIL집단을 2020년에 국립식량과학원 남부작물부 시험포장(경상남도 밀양)에서 적정 이앙기 보다 한달 이른 4월 25일에 조기 재배를 실시하였으며, 15일 동안의 평균기온은 16.8℃, 보통기 이앙을 2020년 5월 30일로 하였을 시, 15일의 평균기온이 23.9℃로 약 7℃정도의 차이를 보였다. 주당 1본씩, 재식거리 30 cm×15 cm로 포장에 이앙하였으며, 농촌진흥청 표준재배법을 준하여 재배하였고, 각 계통 당 초장, 엽수, 건물중을 5반복으로 측정하였다.
Genomic DNA 추출 및 Genotyping
DNA 추출은 파종 후 14일된 유묘의 잎을 채취하여 CTAB (Cetyltrimethyl Ammonium Bromide) 방법으로 추출하였다(Murray & Thompson 1980). 한아름2호×운광 RIL 집단은 IRRI (International Rice Research Institute)의 Genotyping Services Lab.의 BeadXpress (Illumina, San Diego, USA) 384-Plex SNP plates를 이용하여 유전형 데이터를 확보하였다(Thomson et al. 2012). SNP (Single Nucleotide Polymorphism) 마커는 RiceOPA3.1인 Rice SNP set이 사용되었고, Illumina의 GoldenGate Genotyping Assay를 활용하여 Genotyping분석을 진행하였다. BeadXpress Reader (Illumina, San Diego, USA)로 결과를 스캔하고, SNP data는 Illumina Genome eStudio (v 1.1.0)의 Genotyping module로 분석하여 SNP를 calling하였다(Wright et al. 2010).
연관 지도 제작 및 QTL분석
본 연관 지도는 ‘운광’과 ‘한아름2호’사이에 다형성을 보이는 249개의 SNP마커를 이용하여, QTL IciMapping ver. 4.1 (Chinese academy of agricultural sciences, China)로 제작되었다(Meng et al. 2015). 유전적 거리는 Kosambi 함수를 사용하여 추정되었고(Kosambi 1944), 유묘기 저온생장성 연관 QTL은 한아름2호×운광 RIL집단의 표현형 조사 성적을 이용하여 Inclusive composite interval mapping (ICIM-ADD) 방법으로 분석하였다. LOD의 threshold 값은 p<0.05 조건의 1,000 permutation test에 의해 결정되었다(Churchill & Doerge 1994). 후보 유전자 검색은 Rap-db (http://rapdb.dna.affrc.go.jp)을 이용하였다. 형질의 평균 등 기술통계, t-test, Duncan's Multiple Range Test의 통계분석은 R (Version 4.2.1, The R Foundation for Statistical Computing Platform)을 이용하였다.
한아름2호×운광의 RIL집단에 대한 조기 이앙 저온생장성 검정
본 연구에서 조기 이앙 저온생장성 QTL분석을 위하여 조기 이앙 뒤 15일 후 ‘한아름2호’와 ‘운광’, 234개의 RIL집단의 초장, 엽수와 건물중을 측정하였다. 모부본인 ‘한아름2호’와 ‘운광’ 의 초장은 각각 22 cm, 28 cm로 측정되었고, RIL집단의 초장은 17.20-42.87 cm에 분포하였으며, 평균 초장은 28.71 cm였다. 모부본에서 엽수는 각각 5.0개, 3.9개였고, RIL집단 엽수는 3.07-7.00개로 분포하였으며, 평균 엽수는 4.97이였다. 마지막으로 모부본의 건물중은 각각 0.13 g, 0.38 g으로 측정되었고, RIL집단의 건물중은 0.08-1.40 g으로 분포하였으며, 평균 건물중은 0.47 g이였다(Table 1, Figs. 1A, 1B, 1C). RIL집단에 대한 3개의 표현형의 빈도 분포표는 정규분포에 가까운 연속 변이를 나타내었다(Figs. 1A, 1B, 1C). 저온생장성과 관련된 표현형들의 상관분석을 수행한 결과, 고도로 유의한 상관관계를 나타내었다. 상관계수는 초장-건물중(0.78), 건물중-엽수(0.70) 및 초장-엽수(0.46) 순으로 높았다(Fig. 1D).
유전자 지도작성 및 QTL 탐색
한아름2호×운광 RIL집단의 연관 지도는 249개의 SNP로 작성하였으며, 14개의 Linkage group이 만들어졌고, 전체 연관 지도 길이는 1,138.27 cM으로 마커 간 평균 거리는 4.57 cM이었다(Fig. 2). QTL은 주동유전자와 연관된 마커 탐색을 위해서 Permutation test를 1,000회 실행하였으며, 초장, 엽수, 건물중의 Threshold값이 각각 2.83, 2.98, 2.90으로 측정되었다. 각각의 형질에서 해당 Threshold값 이상으로 탐색된 QTL을 확인하였고, 초장에서 1번, 5번 8번, 11번 염색체에서 4개의 QTL (qPH1, qPH5, qPH8, qPH11)이 탐색되었으며, 표현형 변이의 설명력은 5.33~8.45%이었다. 탐색된 QTL의 표현형 변이의 설명력은 qPH8>qPH1>qPH5>qPH11 순으로 높았고, qPH1, qPH5, qPH8의 상가적 효과는 각각 1.13, 0.88, 1.09이며, ‘운광’의 대립유전자형에 의해 초장이 길어지는 효과가 있었다. qPH11의 상가적 효과는 -0.89로 ‘한아름2호’ 대립유전자형에 의해 초장이 길어짐을 알 수 있었다(Table 2, Fig. 3). RIL집단 내에서 ‘한아름2호’ 유래 qPH11만 가지고 있는 그룹은 초장이 27.38 cm로 초장과 연관된 QTL을 지니고 있지 않는 그룹보다 약 2 cm 긴 것으로 확인되었다(Data not shown). qPH11 내에 Glossy1 (GL1), Laccase26 (LAC26) 등의 유전자가 있는 것으로 확인이 되었으며, 초기 생장에 관여하는 유전자로 보고되었다(Islam et al. 2009, Liu et al. 2017). 해당 논문은 통일형 벼의 초기 생장에 저온 피해가 심한 것으로 보고되고 있어, ‘운광’ 유래의 유용 대립유전자의 도입 및 집적 효과 분석이 목적이었으나, ‘한아름2호’ 유래 유용 대립유전자에 대해서는 추가적인 연구가 필요할 것으로 사료된다.
엽수에서는 3번 염색체에서 1개의 QTL (qLN3)이 탐색되었다. qLN3은 LOD값이 4.13이고 표현형 변이의 설명력은 7.56%을 보였다. 상가적 효과가 -0.18이며, ‘한아름2호’ 대립유전자형에 의해 엽수가 증가하는 것으로 판단되었다(Table 2). 통일형 품종은 유묘기에 적고 발생이 심하여 유묘 냉해에 약한 것으로 알려져 있었다(Kim et al. 2017). 반면 본 연구에서 유묘기 저온생장성과 관련된 형질을 세분화하여 분석한 결과, 저온 조건에서 엽수 발생은 내냉성이 강한 자포니카 품종인 ‘운광’에서 유래되지 않고, 통일형 품종인 ‘한아름2호’에서 대립유전자형에서 증가된다는 새로운 사실을 알게 되었다.
건물중에서는 3개의 QTL (qDW1, qDW8, qDW9)이 탐색되었고, 4.52~8.81%의 표현형 변이를 설명하였으며, 3개의 QTL에서 0.05-0.07 g 상가적효과를 나타냈다(Table 2, Fig 3). 각각의 표현형에 대한 QTL 중 초장과 건물중에서 1번과 8번 염색체의 같은 위치에서 QTL이 탐색되었다. qPH1과 qDW1의 QTL 내에서 저온생장성에 연관된 OsTCM1 유전자가 위치하고 있으며, 저온스트레스 내에 엽록체 발달이 조절되는 것으로 보고되었다(Lin et al. 2018). qPH8과 qDW8에서는 OsRH42 유전자가 포함되어 있었으며, pre-mRNA 스플라이싱에 연관된 유전자로 보고되었다. 저온에서 pre-mRNA splicing의 속도가 늦춰지는 것으로 알려져 있으나, OsRH42가 이를 극복하는데 도움을 주는 것으로 보고되었다(Lu et al. 2020). 초장에만 탐색된 qPH5에는 OsLti6 유전자가 보고되었으며, 유묘기 시기에 특정 온도에 원형질막 생성을 조절하여 저온스트레스에 감응하는 유전자로 보고되었다(Morsy et al. 2005). 건물중에만 탐색된 qDW9에는 OsDREB1A와 OsDREB1B가 존재하였으며, DREB (Dehydration-Responsive Element-Binding) 계열에 속하는 유전자이다. 다양한 스트레스에 관여하고 있는 것으로 보고되고 있으며, 저온스트레스뿐만 아니라 내건성, 내염성 등에 관여하는 유전자로 보고되었다(Ito et al. 2005).
QTL 조합에 따른 유전자 집적 효과 분석
통일형 벼 조기이앙 재배 시 저온생장성과 관련된 QTL에 대한 유전자 집적 효과를 분석하였다. 초장과 건물중에서 탐색된 QTL 중에서 ‘운광’의 대립유전자형에 LOD값과 표현형 설명력이 높은 QTL을 선발하였으며, 이들 QTL의 형질별 유전자 집적효과를 분석하였다. ‘운광’ 유래 3개의 초장 QTL에서 QTL이 하나도 집적되지 않은 그룹 1은 25.23 cm로 나타났고, 그룹 2, 3, 4의 qPH1, qPH5, qPH8은 각각 27.63 cm, 27.84 cm, 27.97 cm로 같은 수준의 길이를 보였으며, 하나의 ‘운광’ 유래 QTL을 지니고 있을 때 그룹 1보다 긴 길이를 보였다. 그룹 5의 qPH1과 qPH5의 집적으로 초장의 길이가 28.17 cm로 측정되었지만, ‘운광’ 유래 QTL이 하나만 집적된 그룹들과 통계적으로 같은 수준을 보였으며, qPH5와 qPH8이 집적된 그룹 6과 qPH1과 qPH8이 집적된 그룹 7은 각각 29.88 cm, 30.29 cm로, 그룹 2, 3, 4, 5와 다르지 않은 것으로 확인되었다. ‘운광’ 유래 QTL이 3개가 집적된 그룹 8의 qPH1+qPH5+qPH8이 모든 조합 중 가장 긴 31.72 cm로 측정되었고, 그룹 6의 qPH5+qPH8, 그룹 7의 qPH1+qPH8과 통계적으로 유의한 차이가 없었다(Table 3, Fig. 4A).
‘운광’ 유래의 3개의 건물중 QTL에서 집적되지 않은 그룹 1은 0.37 g으로 가장 가벼운 무게를 보였고, 하나의 ‘운광’ 유래 QTL을 하나만 지닌 그룹 2, 3, 4의 qDW1, qDW8, qDW9는 각각 0.43 g, 0.43 g, 0.44 g로 그룹 1보다 높게 측정되었으나, 통계적으로 비슷한 수준이었다. 두 개의 ‘운광’ 유래 QTL이 집적된 그룹 6의 qDW1+qDW9은 0.57 g, 그룹 5의 qDW1+qDW8은 0.61 g으로 QTL이 하나만 포함된 개체들보다 높은 무게가 측정되었고, 통계적으로 유의한 차이가 있었다. qDW8과 qDW9 조합은 RIL집단에서 탐색 되지 않아 제외되었다. 마지막으로, ‘운광’ 유래 QTL이 3개가 집적된 그룹 7의 qDW1+qDW8+qDW9는 0.63 g으로 그룹들 중 가장 무거운 건물중을 나타냈으나, ‘운광’ 유래 QTL이 2개가 집적된 그룹 5와 그룹 6과는 통계적으로 차이가 없었다(Table 4, Fig. 4B).
엽수에서는 ‘한아름2호’의 대립유전자형으로 단일 QTL만 탐색 되었으며, ‘한아름2호’ 유래 QTL이 없는 그룹 1은 4.76개로 측정되었고 ‘한아름2호’ 유래 QTL이 있는 그룹 2는 5.05개로 두 그룹간 유의하게 차이가 나는 것으로 확인되었다(Table 5).
RIL 집단에서 탐색된 초장과 건물중의 ‘운광’ 유래 QTL과 엽수의 ‘한아름2호’ 유래 QTL을 형질에 상관없이 최대로 집적할 시 5개의 QTL 중 4개의 QTL이 최대로 집적되고, 총 3개의 집적 그룹이 있는 것으로 확인되었다. ‘운광’과 ‘한아름2호’ 유래 5개의 QTL이 집적되지 않은 그룹 1은 초장의 길이가 25.39 cm, 엽수가 4.55 개, 건물중이 0.31 g으로 그룹 중 모든 값이 가장 낮았다. 그룹 2는 초장과 건물중의 QTL 부위가 같은 염색체 1번(qPH1, qDW1)과 엽수의 염색체 3번(qLN3), 초장의 염색체 5번(qPH5), 건물중의 염색체 9번(qDW9)에 있는 QTL들로서 초장, 엽수, 건물중 각각 30.82 cm, 5.67개, 0.63 g으로 측정되었다. 초장과 건물중은 다른 QTL 집적 그룹과 비슷하게 측정되었지만, 엽수는 더 높은 것으로 확인되었다. 그룹 3과 4는 염색체 1번(qPH1, qDW1), 3번(qLN3), 8번(qPH8, qDW8)에 있는 QTL이 공통으로 들어가고, 각각 염색체 5번(qPH5)과 염색체 9번(qDW9)에 있는 QTL이 들어가는 그룹이다. 두 그룹은 초장이 각각 31.24 cm, 31.14 cm로 그룹 2와 비슷한 수준으로 측정되었고, 건물중도 각각 0.53 g, 0.56 g으로 그룹 2와 비슷했지만, 그룹 1과도 차이가 없는 것으로 확인되었다. 엽수는 각각 5.06개, 4.99개로 그룹 2보다는 통계적으로 유의하게 적었다(Table 6). 각각의 형질에서 ‘운광’과 ‘한아름2호’ 유래 QTL이 존재하고 집적될수록 ‘운광’과 ‘한아름2호’ 유래 QTL이 없을 때 보다 해당 표현형이 유의하게 증가하는 것을 확인 할 수 있었다. QTL이 4개 이상 집적된 경우 그룹 2의 조합이 가장 초장이 좋을 것으로 기대되나 표본이 적어 추가적인 연구가 필요할 것으로 사료된다.
유묘기에 저온 피해가 잦은 통일형 품종은 초기 생장성 확보에 어려움을 겪고 있으며, 기후변화로 인한 저온 피해가 증가하고 있는 동아시아에는 저온 적응성을 지닌 품종을 육성하는 것이 요구되고 있다. 통일형 품종의 저온생장성을 향상시키기 위해 통일형 품종인 ‘한아름2호’와 저온생장성이 강한 자포니카 품종 ‘운광’을 교배하여, 234개의 RIL 집단을 육성하고 249개의 SNP마커를 마커를 활용하여 연관 지도 작성 및 QTL분석을 실시하였다. 집단들을 포장에 적정 이앙기보다 한 달 이른 시기에 이앙하여, 초장과 엽수, 건물중을 측정하여 저온생장성에 연관된 QTL을 탐색하였다. 초장에서 저온생장성에 연관된 ‘운광’ 유래 QTL이 각각 염색체 1번, 5번, 8번에서 3개(qPH1, qPH5, qPH8)이 탐색되었고, ‘한아름2호’ 유래 QTL이 염색체 11번에서 1개(qPH11)이 탐색되었다. 엽수에서는 염색체 3번에서 ‘한아름2호’ 유래 QTL 1개(qLN3)만 탐색되었으며, 건물중에서는 ‘운광’ 유래 QTL이 각각 염색체 1번, 8번, 9번에서 3개(qDW1, qDW8, qDW9)이 탐색되었다. 단일 QTL에서 초장은 qPH8이 설명 가능한 표현형이 8.45%, 길이는 27.97 cm로 가장 길었으나 통계적으로 유의한 차이가 없었고, ‘운광’ 유래 3개의 QTL이 집적되었을 시 31.72 cm로 가장 길어졌다. 건물중에서는 ‘운광’ 유래 3개의 QTL이 단일로 각각 존재할 시 통계적으로 차이가 없는 것으로 나왔으며, 3개의 QTL이 집적되었을 시 0.63 g으로 가장 높은 값이 측정되었으나, 2개가 집적된 qDW1+qDW8 (0.61 g), qDW1+qDW9 (0.57 g)과 통계적으로 차이는 없었다. 마지막으로, 엽수의 ‘한아름2호’ 유래 qLN3 여부에 따라 각각 4.76개와 5.05개가 측정되었다. 본 연구에서 탐색된 QTL들을 활용한다면, 저온생장성 통일형 계통 육성이 가능할 것으로 기대한다.
본 논문은 농촌진흥청 연구사업(세부과제명: 대량소비용 산업체 맞춤형 가공용 통일형 벼 육종 소재개발, 세부과제번호: PJ017383012024) 과 2024년도 농촌진흥청 국립식량과학원 전문연구원 과정 지원사업에 의해 이루어진 것이며 이에 감사드립니다.
Fig. 1
Frequency distributions of ‘Hanareum2’ and ‘Unkwang’ recombinant inbred line population at the early planting stage. (A) frequency distributions of plant height in recombinant inbred line population, (B) frequency distributions of leaf number in recombinant inbred line population, (C) frequency distributions of dry weight in recombinant inbred line population, (D) correlation analysis among plant height, leaf number and dry weight. ** significant at p<0.01 (Duncan’s test).
kjbs-56-3-225-f1.jpg
Fig. 2
Linkage maps composed of 249 SNP markers for cold tolerance in 234 recombinant inbred line population. The linkage map was generated using QTL IciMapping program ver. 4.1.
kjbs-56-3-225-f2.jpg
Fig. 3
Quantitative trait loci for plant height, dry weight, and leaf number associated with ‘Unkwang’ alleles were detected on linkage group 1, 5, 9, and 10 in the ‘Hanareum2×Unkwang’ recombinant inbred line population at the early planting stage.
kjbs-56-3-225-f3.jpg
Fig. 4
The quantitative trait loci pyramiding effect in 234 recombinant inbred line populations. (A) Box plot of plant height according to quantitative trait loci. (B) Box plot of dry weight according to quantitative trait loci. Values with a same letter are not significantly different at ** p<0.01 (Duncan’s test).
kjbs-56-3-225-f4.jpg
Table 1
Phenotypic evaluation of plant height, leaf number and dry weight at the early planting stage in recombinant inbred line population.
Table 1
RIL population Traits
Plant height (cm) Leaf number (n) Dry weight (g)
Mean±SD 28.71±3.91 4.97±0.58 0.47±0.21
Range 17.20-42.87 3.07-7.00 0.08-1.40
Table 2
Identification of quantitative trait loci related to plant growth during the early planting stage in a recombinant inbred line population derived from a cross between ‘Hanareum2’ and ‘Unkwang’.
Table 2
QTLs Chromosome
(Linkage group)		 Position
(cM)		 Left marker Right marker LODz PVEy (%) Addx
qPH1 1

(1)		 1

35.39-151.24		 id1018870 id1020384 3
.80		 8

.15		 1

.13
qPH5 5
(6)		 0.00-2.90 id5008218 id5008667 3
.25		 5
.56		 0.88
qPH8 8

(9)		 4.92-25.44 id8000754 id8002025 4.86 8

.45		 1

.09
qPH11 1

1 (12)		 65.17-73.54 id11010555 id11011652 3
.13		 5
.33		 -0.89
qLN3 3
(3)		 1

56.66-159.19		 id3016090 id3016979 4.13 7.56 -0.18
qDW1 1

(1)		 1

35.39-151.24		 id1018870 id1020384 3
.28		 5
.62		 0.06
qDW8 8

(9)		 4.92-25.44 id8000754 id8002025 2.92 4.52 0.05
qDW9 9
(10)		 1

2.95-34.36		 id9006968 id9007180 3
.38		 8

.81		 0.07

zLOD: The logarithm of the odds.

yPVE: Percentage of the phenotypic variance that was explained.

xAdd: Positive value of the additive effect indicate that allele from ‘Unkwang’ was in the direction of increasing the trait value.

Table 3
The quantitative trait loci pyramiding effect of plant height on early planting stage.
Table 3
Group QTL No. of lines Cold tolerance
Plant height (cm)
(Mean±SD)
1 None 32 25.23±4.05cz
2 qPH1 30 27.63±1.96b
3 qPH5 16 27.84±4.05b
4 qPH8 14 27.97±2.70b
5 qPH1+qPH5 18 28.17±2.69b
6 qPH5+qPH8 32 29.88±2.90ab
7 qPH1+qPH8 15 30.29±1.96ab
8 qPH1+qPH5+qPH8 33 31.72±2.86a

zValues with a same letter are not significantly different at **p<0.01 (Duncan’s test).

Table 4
The quantitative trait loci pyramiding effect of dry weight on early planting stage.
Table 4
Group QTL No. of lines Cold tolerance
Plant weight (g)
(Mean±SD)
1 None 32 0.37±0.19bz
2 qDW1 30 0.43±0.14b
3 qDW8 14 0.43±0.12b
4 qDW9 13 0.44±0.11b
5 qDW1+qDW8 15 0.61±0.19a
6 qDW1+qDW9 9 0.57±0.30a
7 qDW1+qDW8+qDW9 8 0.63±0.18a

zValues with a same letter are not significantly different at **p<0.01 (Duncan’s test).

Table 5
The effect of a single quantitative trait locus on leaf number during the early planting stage.
Table 5
Group QTL No. of lines Cold tolerance
Plant weight (g)
(Mean±SD)
1 None 64 4.76±0.54
2 qLN3 154 5.05±0.54**

**p<0.01. Significantly different by Students t-test between non QTL and qLN3.

Table 6
The quantitative trait loci pyramiding effect between plant height, dry weight and leaf number on early planting stage.
Table 6
Group QTL No. of lines Cold tolerance
Plant height (cm) (Mean±SD) Leaf number (n) (Mean±SD) Dry weight (g) (Mean±SD)
1 None 7 25.39±3.92bz 4.55±0.29c 0.31±0.11b
2 qPH1/qDW1+qLN3+qPH5+qDW9 5 30.82±1.63a 5.67±0.59a 0.63±0.13a
3 qPH1/qDW1+qLN3+qPH5+qPH8/qDW8 23 31.24±3.63a 5.06±0.41b 0.53±0.19ab
4 qPH1/qDW1+qLN3+qPH8/qDW8+qDW9 5 31.14±2.58a 4.99±0.20b 0.56±0.12ab

zValues with a same letter are not significantly different at *p<0.01 (Duncan’s test).

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Identification of QTLs Related to Plant Growth at Low Temperatures in the Seedling Stage of Tongil Type Rice after Transplanting
Korean. J. Breed. Sci.. 2024;56(3):225-235.   Published online September 1, 2024
DOI: https://doi.org/10.9787/KJBS.2024.56.3.225
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Identification of QTLs Related to Plant Growth at Low Temperatures in the Seedling Stage of Tongil Type Rice after Transplanting
Korean. J. Breed. Sci.. 2024;56(3):225-235.   Published online September 1, 2024
DOI: https://doi.org/10.9787/KJBS.2024.56.3.225
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Identification of QTLs Related to Plant Growth at Low Temperatures in the Seedling Stage of Tongil Type Rice after Transplanting
Image Image Image Image
Fig. 1 Frequency distributions of ‘Hanareum2’ and ‘Unkwang’ recombinant inbred line population at the early planting stage. (A) frequency distributions of plant height in recombinant inbred line population, (B) frequency distributions of leaf number in recombinant inbred line population, (C) frequency distributions of dry weight in recombinant inbred line population, (D) correlation analysis among plant height, leaf number and dry weight. ** significant at p<0.01 (Duncan’s test).
Fig. 2 Linkage maps composed of 249 SNP markers for cold tolerance in 234 recombinant inbred line population. The linkage map was generated using QTL IciMapping program ver. 4.1.
Fig. 3 Quantitative trait loci for plant height, dry weight, and leaf number associated with ‘Unkwang’ alleles were detected on linkage group 1, 5, 9, and 10 in the ‘Hanareum2×Unkwang’ recombinant inbred line population at the early planting stage.
Fig. 4 The quantitative trait loci pyramiding effect in 234 recombinant inbred line populations. (A) Box plot of plant height according to quantitative trait loci. (B) Box plot of dry weight according to quantitative trait loci. Values with a same letter are not significantly different at ** p<0.01 (Duncan’s test).

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Identification of QTLs Related to Plant Growth at Low Temperatures in the Seedling Stage of Tongil Type Rice after Transplanting

Phenotypic evaluation of plant height, leaf number and dry weight at the early planting stage in recombinant inbred line population.

RIL population Traits
Plant height (cm) Leaf number (n) Dry weight (g)
Mean±SD 28.71±3.91 4.97±0.58 0.47±0.21
Range 17.20-42.87 3.07-7.00 0.08-1.40

Identification of quantitative trait loci related to plant growth during the early planting stage in a recombinant inbred line population derived from a cross between ‘Hanareum2’ and ‘Unkwang’.

QTLs Chromosome
(Linkage group)		 Position
(cM)		 Left marker Right marker LODz PVEy (%) Addx
qPH1 1

(1)		 1

35.39-151.24		 id1018870 id1020384 3
.80		 8

.15		 1

.13
qPH5 5
(6)		 0.00-2.90 id5008218 id5008667 3
.25		 5
.56		 0.88
qPH8 8

(9)		 4.92-25.44 id8000754 id8002025 4.86 8

.45		 1

.09
qPH11 1

1 (12)		 65.17-73.54 id11010555 id11011652 3
.13		 5
.33		 -0.89
qLN3 3
(3)		 1

56.66-159.19		 id3016090 id3016979 4.13 7.56 -0.18
qDW1 1

(1)		 1

35.39-151.24		 id1018870 id1020384 3
.28		 5
.62		 0.06
qDW8 8

(9)		 4.92-25.44 id8000754 id8002025 2.92 4.52 0.05
qDW9 9
(10)		 1

2.95-34.36		 id9006968 id9007180 3
.38		 8

.81		 0.07

The quantitative trait loci pyramiding effect of plant height on early planting stage.

Group QTL No. of lines Cold tolerance
Plant height (cm)
(Mean±SD)
1 None 32 25.23±4.05cz
2 qPH1 30 27.63±1.96b
3 qPH5 16 27.84±4.05b
4 qPH8 14 27.97±2.70b
5 qPH1+qPH5 18 28.17±2.69b
6 qPH5+qPH8 32 29.88±2.90ab
7 qPH1+qPH8 15 30.29±1.96ab
8 qPH1+qPH5+qPH8 33 31.72±2.86a

The quantitative trait loci pyramiding effect of dry weight on early planting stage.

Group QTL No. of lines Cold tolerance
Plant weight (g)
(Mean±SD)
1 None 32 0.37±0.19bz
2 qDW1 30 0.43±0.14b
3 qDW8 14 0.43±0.12b
4 qDW9 13 0.44±0.11b
5 qDW1+qDW8 15 0.61±0.19a
6 qDW1+qDW9 9 0.57±0.30a
7 qDW1+qDW8+qDW9 8 0.63±0.18a

The effect of a single quantitative trait locus on leaf number during the early planting stage.

Group QTL No. of lines Cold tolerance
Plant weight (g)
(Mean±SD)
1 None 64 4.76±0.54
2 qLN3 154 5.05±0.54**

The quantitative trait loci pyramiding effect between plant height, dry weight and leaf number on early planting stage.

Group QTL No. of lines Cold tolerance
Plant height (cm) (Mean±SD) Leaf number (n) (Mean±SD) Dry weight (g) (Mean±SD)
1 None 7 25.39±3.92bz 4.55±0.29c 0.31±0.11b
2 qPH1/qDW1+qLN3+qPH5+qDW9 5 30.82±1.63a 5.67±0.59a 0.63±0.13a
3 qPH1/qDW1+qLN3+qPH5+qPH8/qDW8 23 31.24±3.63a 5.06±0.41b 0.53±0.19ab
4 qPH1/qDW1+qLN3+qPH8/qDW8+qDW9 5 31.14±2.58a 4.99±0.20b 0.56±0.12ab

zValues with a same letter are not significantly different at *p<0.01 (Duncan’s test).

Table 1 Phenotypic evaluation of plant height, leaf number and dry weight at the early planting stage in recombinant inbred line population.
Table 2 Identification of quantitative trait loci related to plant growth during the early planting stage in a recombinant inbred line population derived from a cross between ‘Hanareum2’ and ‘Unkwang’.

zLOD: The logarithm of the odds.

yPVE: Percentage of the phenotypic variance that was explained.

xAdd: Positive value of the additive effect indicate that allele from ‘Unkwang’ was in the direction of increasing the trait value.

Table 3 The quantitative trait loci pyramiding effect of plant height on early planting stage.

zValues with a same letter are not significantly different at **p<0.01 (Duncan’s test).

Table 4 The quantitative trait loci pyramiding effect of dry weight on early planting stage.

zValues with a same letter are not significantly different at **p<0.01 (Duncan’s test).

Table 5 The effect of a single quantitative trait locus on leaf number during the early planting stage.

**p<0.01. Significantly different by Students t-test between non QTL and qLN3.

Table 6 The quantitative trait loci pyramiding effect between plant height, dry weight and leaf number on early planting stage.

zValues with a same letter are not significantly different at *p<0.01 (Duncan’s test).

Table 1

Table 2

Table 3

Table 4

Table 5

Table 6

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