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바스마티 370 품종을 활용한 벼 종자 크기 관련 양적형질 유전자좌 분석

임다은1, 장성규1,2, 김백기1,2, 서정환3,4, 키쇼르3, 고희종3, 권순욱1,2,*

QTL Analysis Related to Grain Size Using the Population Derived from a Cross Between Hopum and Basmati 370

Da-Eun Im1, Seong-Gyu Jang1,2, Backki Kim1,2, Jeonghwan Seo3,4, D. S. Kishor3, Hee-Jong Koh3, Soon-Wook Kwon1,2,*
Korean Journal of Breeding Science 2023;55(2):118-125.
Published online: June 1, 2023

1부산대학교 식물생명과학과

2부산대학교 생명산업융합연구원

3서울대학교 농림생물자원과학부

4농촌진흥청 국립식량과학원 작물육종과

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*Corresponding Author (E-mail: swkwon@pusan.ac.kr, Tel: +82-55-350-5506, Fax: +82-55-350-5509)
• Received: May 18, 2023   • Revised: May 18, 2023   • Accepted: May 23, 2023

Copyright © 2023 by the Korean Society of Breeding Science

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  • The grain size and shape have a significant impact on the grain quality and yield of rice. Additionally, due to the globalization of the rice market, there is a demand for rice varieties of various sizes. In this study, we analyzed quantitative trait loci (QTL) associated with grain length (GL), grain width (GW), grain thickness (GT) and ratio of length-to-width (RLW) using 149 F2 plants obtained from a cross between Hopum and Basmati 370. To construct a linkage map, we utilized 89 Fluidigm SNP markers, which covered a span of 1,116 cM of F2 population. Nine QTLs associated with four traits were identified on chromosomes 2, 3, 5, and 7. One major QTL on chromosome 3 accounted for 43.2% of the phenotypic variation in GL. Additionally, QTLs named qGW2 and qGW5 on chromosomes 2 and 5 were detected for GW, explaining 10.4% and 18.6% of the phenotypic variation, respectively. Three QTLs for GT were found on chromosomes 2, 5, and 7, explaining 9.9%, 12.9%, and 9.9% of phenotypic variation, respectively. These QTLs offer potential applications in breeding programs using Basmati 370 to develop long and slender grain japonica varieties.
벼는 세계에서 가장 중요한 식량 작물 중 하나이며 세계인구 절반의 주식으로 이용되고 있다(Zeng et al. 2017). 벼의 수량성 향상은 오랜 시간동안 가장 중요한 육종 목표 중 하나로, 주당수수, 수당립수, 종자무게의 영향을 가장 크게 받는다. 종자무게는 종자 크기와 정의 상관으로 연관되어있다. 따라서 종자 크기는 벼의 수확량을 향상시키는 중요한 농업 특성이다. 더욱이 벼의 종자 크기와 형태는 수량과 품질에 있어 중요한 역할을 하는 형질이다(Harberd 2015, Xing & Zhang 2010). 벼의 종자 크기는 종자 길이(grain length, GL), 종자 폭(grain width, GW), 종자 두께(grain thickness, GT)와 장폭비(ratio of length to width, RLW)에 의해 결정된다(Zuo & Li 2014). 재배벼의 생태형은 주로 인디카와 자포니카로 구분하는데, 종자 길이가 길고 종자 폭이 좁은 장립종의 인디카는 세장형 입형을 주로 보이며, 그에 비해 자포니카는 종자 길이가 짧고 형태가 둥근 단원형 입형을 주로 보인다(Juliano 1979, Kang et al. 2006).
우리나라를 포함한 동북아시아 지역에서는 쌀이 짧고 둥근 형태를 선호하는 반면, 동남아시아 및 서구 지역은 길고 얇은 형태의 쌀을 선호하는 것으로 알려져 있다(Calingacion et al. 2014). 특히 우리나라의 경우 찰기가 많고 끈기가 있는 자포니카형 벼의 선호도가 특히 높아서, 인디카와 자포니카 교잡을 통해 개발된 통일형 품종들까지도 단원형 입형을 보이게 육성하고자 하였다. 따라서 우리나라 품종들은 대부분 중단립종 단원형을 모양을 보이고 있고, 유전전 배경이 협소해짐에 따라 새롭게 개발되는 품종들의 경우에도 입형은 중단립종 단원형 위주로 고착화되었다(Park et al. 2017). 대립벼 1호(Shim et al. 2016), 대립자미벼(Han et al. 2012), 슈퍼자미 2호(Ham et al. 2015) 등의 품종은 우리나라에서 종자 크기가 큰 대립종 품종이지만, 장폭비는 자포니카형에 가까워 입형다양화에 대한 요구는 충족시키지 못하고 있는 실정이다. 국내 기후변화에 따른 품질저하 문제를 극복하고, 쌀 수출 확대를 위한 전략을 세우기 위해 국내 벼 품종의 입형다양화에 대한 요구는 커지고 있다(Kim et al. 2020).
종자 크기에 관련된 형질들은 여러 유전자에 의해 조절되는 양적형질로 알려져 있으며 지금까지 양적형질유전자좌(quantitative trait loci; QTL)가 400여개 이상 탐색 되었으며, 일부는 유전자 수준에서 기능이 규명되었다(Huang et al. 2013, Zhou et al. 2015). 종자 크기와 관련하여 최초로 cloning 된 Grain size 3 (GS3) 유전자는 종자 길이와 무게에 주동으로 작용하는 유전자로 기관 크기 조절(organ size regulation; ORS) 도메인 기능으로 종자 길이를 조절하는 것으로 보고되었다(Fan et al. 2006). qGL3.1 (QTL for grain length 3.1)은 단백질 인산화에 영향을 주어 종자 길이와 수량을 조절한다(Qi et al. 2012). Grain width and weight 2 (GW2) 유전자는 종자 폭에 영향을 준다고 보고되어 있는데, 유전자의 기능이 상실되었을 때 세포 수를 증가시켜 종자 폭 및 무게를 증가시킨다고 알려져 있다(Song et al. 2007). qSW5/GW5 (QTL for seed width 5/grain width 5) 유전자는 종자의 폭과 무게를 조절한다고 보고되어 있는데, 유전자의 1212 bp가 결실되었을 때 종자의 폭과 무게를 증가되는 것으로 알려져 있다(Shomura et al. 2008, Weng et al. 2008). Grain size 5 (GS5) 유전자는 qSW5/GW5 유전자와 2 Mb 거리에 연관되어 있고, 유전자 발현이 활성화되었을 때 종자 크기를 증가시키는 것으로 알려져 있다(Li et al. 2011). 그 외에도 GW8 (Wang et al. 2012), TGW6 (Ishimaru et al. 2013), GW7 (Wang et al. 2015), GLW7 (Si et al. 2016), LGY3 (Liu et al. 2018) 등의 유전자가 종자 크기와 관련된 기능에 대해 연구되었다. 특히 입형다형화를 위해 야생벼(O. grandiglumis, O. rufipogon) 유래의 종자 크기 연관 유전자 연구도 진행된 바 있다(Kang et al. 2016, Kim et al. 2015).
본 연구는 국내 품종의 종자 크기 다양화를 위해 ‘호품’과 ‘Basmati 370’을 교배한 F2집단에 대해 종자 크기를 평가하고, 총 89개의 SNP마커를 이용하여 종자 크기 관련 QTL 탐색하기 위해 수행하였다.
시험 재료 및 재배방법
본 연구에서 종자크기 관련 QTL 분석을 위하여 ‘호품’과 ‘Basmati 370’의 인공교배를 통해 육성된 149개의 F2집단을 이용하였다. ‘호품’은 직파 적응성 최고품질 내도복 자포니카 품종(Ko et al. 2008)으로 종자 길이, 종자 너비는 중간 정도이며, 현미모양은 단원형이다. ‘Basmati 370’은 아시아뿐만 아니라 서구에서도 인기가 있는 aromatic형 향미 품종(Bhattacharjee et al. 2002, Kishor et al. 2020)으로 종자 길이는 장립종 정도로 길고, 현미 모양은 세장형이다.
종자크기 특성 조사
표현형 조사는 종자 크기에 연관된 세가지 형질을 수확된 종자를 이용하여 수행하였다. 수확한 종자는 형질을 조사하기 전에 통풍이 잘되는 상온에서 건조하고 보관하였다. 충실한 종자 중 무작위로 선택한 종자 10개의 종자 길이, 폭, 두께를 측정하여 각 식물체 별로 평균을 산출하였다. 세 가지 형질은 버니어 캘리퍼스를 이용하여 조사되었다. 장폭비는 종자 길이와 종자 폭의 비율로 계산하였다.
DNA 추출
DNA는 벼의 어린 잎을 채취하여, CTAB (cetyltrimethyl ammonium bromide) 방법(Murray & Thompson 1980)을 응용하여 추출하였다. 잎 1 g을 막자사발에 놓고 액체 질소에 침적시킨 후 마쇄하였다. 마쇄된 잎 샘플에 600 μL의 CTAB solution을 넣고, 65℃로 30분 처리하였다. 600 μL CI 용액(chloroform: isoamyl alcohol=24:1)을 첨가한 후 12,000 rpm으로 5분간 원심분리하여 400 μL 상등액을 1.5 mL tube에 옮겼다. 400 μL의 isopropyl alcohol을 첨가하여 2~3회 혼합한 후 -20℃로 3시간 처리한 뒤 12,000rpm에서 10분간 원심분리 하였다. 상등액을 버리고 침전물을 70% ethanol로 2회 세정 후 건조하였으며, TE buffer (10 mM Tris-HCL, ph 8.0, 2.5 mM EDTA)에 녹인 후 유전자형 분석에 이용했다.
Fluidigm 유전자형 분석
유전자형 분석을 위하여 기존에 보고되어 있는 총 192개의 indica-japonica Fluidigm SNP 마커를 이용하여 분석하였다(Seo et al. 2020). ‘호품’과 ‘Basmati 370’ 교배를 통해 육성된 F2 집단의 종자 크기 관련 3가지 형질과 연관된 QTL을 결정하기 위해 사용되었다. Fluidigm SNP 유전자형 분석은 그린바이오 공동기기센터 NICEM 평창분원에서 BioMarkTMHD system (Fluidigm, San Francisco, CA, USA), 96.96 Dynamic Array IFCs를 사용하여 수행되었다. Fluidigm SNP 유전자형 분석 프로그램을 이용하여 유전자형 결과를 얻고, 모든 유전자형을 수동으로 추가 확인하여 엄격히 평가하였다.
연관지도(linkage map) 작성 및 QTL 분석
연관지도는 총 192개의 Fluidigm SNP 마커 중 ‘호품’과 ‘Basmati 370’에서 다형성을 보인 89개의 마커에 대해 IciMapping ver 4.1 (Chinese Acadeny of agrocultural Sciencs, China)를 이용하여 작성하였다(Meng et al. 2015). QTL 분석은 ICIM (Inclusive composite interval mapping) 방법으로 수행하였고, 유의한 QTL을 결정하기 위한 LOD의 임계값은 p≤0.05 조건으로 1,000반복한 permutation test에 의해 결정하였다. 후보 유전자는 Rap-db (https://rapdb.dna.affrc.go.jp/)를 이용하여 검색하였다. 형질의 평균 등 기술통계, t-test, Duncan’ Multiple Range Test의 통계분석은 R (Version 4.3.0, The R Foundation for Statistical Computing Platform)을 사용하였다.
표현형 조사와 연관지도 작성
종자 크기와 관련된 QTL부위를 탐색하기 위해 두 모본과 149개 F2에 종자 길이, 종자 폭, 종자 두께 및 장폭비를 조사하였다. ‘호품’의 각 표현형은 순서대로 7.25 mm, 3.50 mm, 2.17 mm, 2.07 이었고, ‘Basmati 370’은 10.17 mm, 2.56 mm, 1.97 mm, 3.97이었다.
F2집단에서 각각 종자 길이, 종자 폭, 종자 두께 및 장폭비는 6.59~9.18 mm, 2.06~3.25 mm, 1.50~2.17 mm, 2.36~4.19로 나타났으며 평균 길이, 폭, 두께 및 장폭비는 각각 7.88 mm, 2.57 mm, 1.88 mm, 3.09으로 나타났다(Fig. 1).
F2 집단에서 4가지 표현형 간의 관계를 확인하기 위해 상관분석을 수행하였다. 표현형 간의 상관관계는 Table 1과 같다. 종자 폭과 두께 사이에서 고도로 유의한 양의 상관관계(0.60)를 보였고, 종자 폭과 장폭비 사이에서 고도로 유의한 부의 상관관계(-0.79)를 보였다. 장폭비는 종자의 길이, 폭, 두께와 모두 유의한 정의상관 또는 부의상관을 보였다. 이는 4가지 특성이 동일하거나 밀접하게 연관된 QTL에 의해 조절될 수 있음을 나타낸다.
‘호품’과 ‘Basmati 370’에서 다형성을 보이는 총 89개의 마커를 선발하였고, 해당 마커의 SNP정보를 이용하여 F2집단의 연관지도를 작성하였다(Fig. 2). 염색체별로 3~14개 마커가 배치되었다. 연관지도의 총 길이는 1,116 cM이고 마커간 평균거리는 14.50 cM이다. 대부분의 염색체에서 마커간 간격이 긴 구간이 존재하거나 연관지도가 전체 염색체를 포함하지 못하였다. 이는 본 연구에서 사용한 SNP 마커 세트가 인디카와 자포니카 다형성을 구분하기 위한 마커 세트이기 때문이다. ‘호품’과 ‘Basmati 370’는 190개 indica-japonica SNP를 이용한 유전다양성 분석에서 각각 온대 자포니카와 aromatic 그룹에 속하였으며, Aromatic 그룹은 인디카보다 자포니카 그룹에 더 가까운 유연관계를 보였다(Kishor et al. 2020). 향후 연관지도의 공백을 채우기 위해서는 ‘호품’과 ‘Basmati 370’ 간의 다형성 SNP 추가가 필요하다.
QTL 분석
‘호품’과 ‘Basmati 370’의 종자크기를 제어하는 유전자 영역을 탐색하기 위해 종자 길이, 종자 폭, 종자 두께 및 장폭비 표현형에 대한 permutation test로 얻어진 임계 LOD값 3.96, 3.31, 3.52, 4.08 이상인 QTL을 선발하였다. 각 표현형 별로 선발된 QTL은 Table 2와 같다.
염색체 3번에서 탐색된 qGL3과 qRLW3은 GS3-CA와 id3010700사이에 위치하였다. qGL3의 LOD는 20.1이며 GL 표현형 변이의 43.15%를 설명하였고, qRLW3의 LOD는 10.4이며 RLW의 표현형 변이의 20.36%를 설명하였다. 이 영역에는 종자 길이와 무게에 주동으로 작용하는 유전자로 알려진 GS3 유전자(Chr. 3: 16,729,501-16,735,109, https://rapdb.dna.affrc.go.jp/)가 존재하고 있으며, 종자 길이에 부의 조절 작용(negatively regulator)을 하는 것으로 알려져 있다. 이 유전자의 두 번째 exon 영역에서 발생하는 C→A로의 염기치환은 translation stop codon (TGA)을 생성하여 nonsense mutation을 일으키게 되고, 유전자의 기능을 상실시켜 종자 길이를 증가시킨다고 보고되었다(Fan et al. 2009). PstⅠ 제한효소를 이용하여 C→A 염기치환을 검정할 수 있는 cleaved amplified polymorphic sequence (CAPS) 마커가 개발되었다(Yan et al. 2009).
염색체 5번에서 탐색된 qSW5와 qRLW5는 cmb0501.9와 id5002497사이에 위치하고, qGT5는 id5002497와 qSW5-AG 사이에 위치하고 있다. qSW5의 LOD는 8.5이며 전체 종자 폭의 표현형 변이 중 18.60%를 설명하였고, qRLW5의 LOD는 7.1으로 장폭비의 표현형 변이를 15.13%를 설명하였다. qGT5의 LOD는 6.0이며 종자 두께 표현형 변이의 12.88%를 설명하였다. 이 영역에는 5번 염색체에서 GW와 무게에 관여하는 주동 QTL로 알려진 qSW5/GW5가 존재하고 있다. 이 유전자 염기서열 1212 bp 결실은 대부분의 자포니카 벼에서 발견되는데 이는 유전자 기능을 상실시켜 종자 폭은 넓어지고 무게는 증가하게 된다(Shomura et al. 2008). 발견된 QTL 영역에 보고 되어있는 GS3과 qSW5/GW5 간에 종자의 길이와 폭 조절에 있어 상호작용이 있다고 보고되었다(Yan et al. 2011).
종자 길이, 종자 폭, 종자 두께 및 장폭비별 탐색된 QTL의 표현형 변이 설명 정도는 각각 43.15%, 28.96%, 32.68%, 45.38%로서 다소 낮았는데, 이는 연관지도가 전체 염색체 영역을 포함하지 못했기 때문으로 보인다. 다만 입형관련 주동유전자에 대해 GW2, GS3, GW5에 의한 것이라는 기존 보고와 일치한다(Park et al. 2017).
QTL 집적에 따른 종자크기 차이 분석
종자 크기와 관련하여 탐색된 QTL의 집적 효과를 검토하고자 QTL영역으로 탐색된 부분에 위치한 연관 마커를 활용하여 QTL의 집적 여부에 따라 호품/Basmati 370 F2집단을 각 2~8개의 그룹으로 나누었다(Table 3, Fig. 3).
종자 길이는 qGL3이 탐색되지 않은 그룹 1은 7.726 mm 표현형을 보였고, qGL3를 포함한 그룹 2의 종자 길이는 8.486 mm로 qGL3의 도입에 의한 종자 길이의 증가가 뚜렷하게 나타났다.
종자 폭 관련 QTL이 집적 여부에 따라 4개 그룹으로 구분되었고, QTL이 집적되지 않은 그룹 1은 2.510 mm 표현형을 보였고, qGW2이 도입된 그룹 2는 2.613 mm이고, qGW5이 도입된 그룹 3은 2.765 mm 로 그룹 1보다 넓어졌다. qGW2과 qGW5이 집적된 그룹 4의 종자 폭은 2.845 mm 로 가장 넓은 표현형을 나타내는 것으로 조사되었다. 두 QTL 사이에 상가적 효과가 있는 것으로 보였다.
종자 두께는 3개의 QTL이 탐색되었고, 이들의 집적 여부에 따라 7개 그룹으로 구분되었다. QTL이 집적되지 않은 그룹 1은 1.836 mm 표현형을 보였고, qGT2이 도입된 그룹 2는 1.889 mm로, qGT5이 도입된 그룹 3은 1.917 mm 와 qGT7은이 도입된 그룹 4는 1.922mm로 비슷한 수준을 보였다. 2개의 QTL이 집적된 그룹 5, 그룹 6, 그룹 7은 각각 2.043 mm, 1.983 mm, 2.034 mm로 두꺼워 졌다.
장폭비는 종자 길이와 높은 정의상관을 보이고 종자 폭과는 부의 상관을 보였다(Table 1). 따라서 장폭비 관련 QTL은 qGL3, qGW2, qGW5와 동일한 영역에서 탐색되었다. QTL이 집적되지 않은 그룹 1은 2.925의 수준을 보였고, qRLW2, qRLW3, qRLW5가 각각 하나씩 도입된 그룹 2, 3, 4의 장폭비는 각각 3.128, 3.260, 3.168로 나타났다. 2개의 QTL이 집적된 그룹 5 (3.649), 그룹 6 (3.299), 그룹 7 (3.612)에서 증가하였고, 3개의 QTL이 집적된 그룹 8 (3.582)은 그룹 7(3.612)과 비슷한 값을 나타냈다. QTL이 단독으로 집적되었을 때는 qRLW3이 qRLW2와 qRLW5보다 장폭비 증가에 더 효과적이었고, qRLW2+qRLW3 조합과 qRLW3+qRLW5 조합에서 집적효과가 큰 것으로 나타났다. 이는 종자 길이의 변이가 종자 폭의 변이에 비해 크게 나타났고, 종자 길이와 폭과 관련된 QTL이 동시에 집적된 경우 장폭비를 가장 크게 증가시키는 것으로 나타났다(Table 3).
본 연구는 호품/Basmati 370 F2집단을 이용하여 종자 크기에 관련된 형질들에 대해 QTL분석을 수행하였다. 유전자 연관지도는 두 모본에서 다형성을 보인 89개의 SNP 마커를 이용하여 작성되었다. 종자 길이와 관련된 QTL이 1개, 종자 폭에 관련된 QTL이 2개, 종자 두께에 관련된 QTL이 3개, 장폭비와 관련된 3개 QTL이 탐색되었다.
종자 길이와 관련된 qGL3는 표현형 변이를 43.15% 설명하는 주동유전자 부위로 ‘Basmati 370’에서 유래되었고, 그 위치에는 GS3가 보고된 바 있다. 종자 폭과 관련된 2개의 QTL (qGW2, qGW5)이 탐색되었고, 각각 표현형 변이를 10.36%, 18.60%를 설명하는 QTL 영역으로 특히 qGW5 영역에는 qSW5가 보고된 바 있다. 종자 두께와 관련된 3개의 QTL (qGT2, qGT5, qGT7)이 탐색되었고, 각각 표현형 변이의 9.88%, 12.88%, 9.92%를 설명하였으며, ‘호품’에서 유래되었다. 장폭비와 관련된 3개의 QTL (qRLW2, qRLW3, qRLW5)은 종자 길이와 종자 폭과 관련된 QTL과 동일한 영역에서 탐색되었으며 이들의 표현형 변이값은 각각 9.89, 20.36, 15.13% 이였으며, 전체 장폭비 변이의 45.38%를 포함하였다.
본 연구는 한국연구재단 이공학개인기초연구사업(과제번호: 2018R1D1A1B07051390)과와 농촌진흥청 공동연구사업(과제번호: PJ01707303 )에 의해 이루어진 것입니다.
Fig. 1
Phenotypic distribution of traits related to grain size using the 149 F2:3 lines derived from a cross between Hopum and Basmati 370. GL: grain length, GW: grain width, GT: grain thickness, RLW: ratio of length to width.
KJBS-55-118-f1.tif
Fig. 2
Linkage maps composed of 89 indica-japonica Fluidigm SNP markers for grian size related traits in 149 F2 population drived from a cross between Hopum and Basmati 370. Grain length (GL) related QTL was detected on chromosome 3. Grain width (GW) related QTLs were detected on chromosome 2, 5. Grain thickness (GT) related QTLs were detected on chromosome 2, 5 and 7. The ratio of length to width (RLW) related QTLs were detected on chromosome 2, 3 and 5. The linkage map was generated using QTL IciMapping program ver. 4.1. Red : GL, Green: GW, Mint: GT, Blue: RLW.
KJBS-55-118-f2.tif
Fig. 3
The Grain size related traits of the Hopum/Basmati 370 F2 carrying different allele combination of each QTLs. Letters from a to d indicate significantly different values according to Duncan's multiple range test (α=0.05). Errors bars indicate the standard errors of each category.
KJBS-55-118-f3.tif
Table 1
Correlation coefficients among the GL, GW, GT, and RLW in the F2 population.
Table 1
Traitsz GL GW GT RLW
GL 1
GW 0.02 1
GT 0.16 0.6** 1
RLW 0.58** -0.79** -0.39** 1

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

**Indicate significant at p=0.01 level, respectively.

Table 2
QTL for grain size related traits detected in the F2 population derived from a cross between Hopum and Basmati 370.
Table 2
Traitsz QTL Chr. Position (cM) Left Marker Right Marker LODy PVEx (%) Addw Reported Gene/QTL
GL qGL3 3 76 GS3-CA id3010700 20.1 43.15 -0.4611 GS3
GW qGW2 2 79 cmb0219.1 ah02001499 4.9 10.36 0.1022
qGW5 5 18 cmb0501.9 id5002497 8.5 18.60 0.1581 qSW5
GT qGT2 2 86 ah02001499 id2009889 4.4 9.88 0.054
qGT5 5 22 id5002497 qSW5-AG 6.0 12.88 0.0639 qSW5
qGT7 7 81 id7003072 SLG7-GC 4.6 9.92 0.049
RLW qRLW2 2 81 cmb0219.1 ah02001499 5.3 9.89 -0.1375
qRLW3 3 76 GS3-CA id3010700 10.4 20.36 -0.2221 GS3
qRLW5 5 16 cmb0501.9 id5002497 7.1 15.13 -0.1989 qSW5

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

yLOD: The logarithm of the odds.

xPVE: Percentage of the phenotypic variance that was explained.

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

Table 3
QTL pyramiding effect on grain size related traits. Letters from a to d indicate significantly different values according to Duncan's multiple range test (α=0.05).
Table 3
Traitsz group QTL No. of lines Range (Mean±SD)
GL
(mm)		 1 Non 118 7.726±0.423b
2 qGL3 31 8.486±0.431a
GW
(mm)		 1 Non 99 2.510±0.224c
2 qGW2 24 2.613±0.158bc
3 qGW5 23 2.765±0.240ab
4 qGW2+qGW5 3 2.845±0.038a
GT
(mm)		 1 Non 82 1.836±0.117d
2 qGT2 20 1.889±0.078cd
3 qGT5 23 1.917±0.096bcd
4 qGT7 12 1.922±0.088bcd
5 qGT2+qGT5 4 2.043±0.019a
6 qGT2+qGT7 4 1.983±0.098abc
7 qGT5+qGT7 4 2.034±0.099abc
8 qGT2+qGT5+qGT7 0 -
RLW 1 Non 75 2.925±0.251d
2 qRLW2 17 3.128±0.342cd
3 qRLW3 16 3.260±0.304bcd
4 qRLW5 26 3.168±0.333cd
5 qRLW2+qRLW3 4 3.649±0.167a
6 qRLW2+qRLW5 3 3.299±0.159abc
7 qRLW3+qRLW5 5 3.612±0.471ab
8 qRLW2+qRLW3+qRLW5 3 3.582±0.512ab

Means with same letters are not significantly different at p<0.05 (ANOVA followed by DMRT).

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

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QTL Analysis Related to Grain Size Using the Population Derived from a Cross Between Hopum and Basmati 370
Korean. J. Breed. Sci.. 2023;55(2):118-125.   Published online June 1, 2023
DOI: https://doi.org/10.9787/KJBS.2023.55.2.118
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QTL Analysis Related to Grain Size Using the Population Derived from a Cross Between Hopum and Basmati 370
Korean. J. Breed. Sci.. 2023;55(2):118-125.   Published online June 1, 2023
DOI: https://doi.org/10.9787/KJBS.2023.55.2.118
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QTL Analysis Related to Grain Size Using the Population Derived from a Cross Between Hopum and Basmati 370
Image Image Image
Fig. 1 Phenotypic distribution of traits related to grain size using the 149 F2:3 lines derived from a cross between Hopum and Basmati 370. GL: grain length, GW: grain width, GT: grain thickness, RLW: ratio of length to width.
Fig. 2 Linkage maps composed of 89 indica-japonica Fluidigm SNP markers for grian size related traits in 149 F2 population drived from a cross between Hopum and Basmati 370. Grain length (GL) related QTL was detected on chromosome 3. Grain width (GW) related QTLs were detected on chromosome 2, 5. Grain thickness (GT) related QTLs were detected on chromosome 2, 5 and 7. The ratio of length to width (RLW) related QTLs were detected on chromosome 2, 3 and 5. The linkage map was generated using QTL IciMapping program ver. 4.1. Red : GL, Green: GW, Mint: GT, Blue: RLW.
Fig. 3 The Grain size related traits of the Hopum/Basmati 370 F2 carrying different allele combination of each QTLs. Letters from a to d indicate significantly different values according to Duncan's multiple range test (α=0.05). Errors bars indicate the standard errors of each category.

Fig. 1

Fig. 2

Fig. 3

QTL Analysis Related to Grain Size Using the Population Derived from a Cross Between Hopum and Basmati 370

Correlation coefficients among the GL, GW, GT, and RLW in the F2 population.

Traitsz GL GW GT RLW
GL 1
GW 0.02 1
GT 0.16 0.6** 1
RLW 0.58** -0.79** -0.39** 1

QTL for grain size related traits detected in the F2 population derived from a cross between Hopum and Basmati 370.

Traitsz QTL Chr. Position (cM) Left Marker Right Marker LODy PVEx (%) Addw Reported Gene/QTL
GL qGL3 3 76 GS3-CA id3010700 20.1 43.15 -0.4611 GS3
GW qGW2 2 79 cmb0219.1 ah02001499 4.9 10.36 0.1022
qGW5 5 18 cmb0501.9 id5002497 8.5 18.60 0.1581 qSW5
GT qGT2 2 86 ah02001499 id2009889 4.4 9.88 0.054
qGT5 5 22 id5002497 qSW5-AG 6.0 12.88 0.0639 qSW5
qGT7 7 81 id7003072 SLG7-GC 4.6 9.92 0.049
RLW qRLW2 2 81 cmb0219.1 ah02001499 5.3 9.89 -0.1375
qRLW3 3 76 GS3-CA id3010700 10.4 20.36 -0.2221 GS3
qRLW5 5 16 cmb0501.9 id5002497 7.1 15.13 -0.1989 qSW5

QTL pyramiding effect on grain size related traits. Letters from a to d indicate significantly different values according to Duncan's multiple range test (α=0.05).

Traitsz group QTL No. of lines Range (Mean±SD)
GL
(mm)		 1 Non 118 7.726±0.423b
2 qGL3 31 8.486±0.431a
GW
(mm)		 1 Non 99 2.510±0.224c
2 qGW2 24 2.613±0.158bc
3 qGW5 23 2.765±0.240ab
4 qGW2+qGW5 3 2.845±0.038a
GT
(mm)		 1 Non 82 1.836±0.117d
2 qGT2 20 1.889±0.078cd
3 qGT5 23 1.917±0.096bcd
4 qGT7 12 1.922±0.088bcd
5 qGT2+qGT5 4 2.043±0.019a
6 qGT2+qGT7 4 1.983±0.098abc
7 qGT5+qGT7 4 2.034±0.099abc
8 qGT2+qGT5+qGT7 0 -
RLW 1 Non 75 2.925±0.251d
2 qRLW2 17 3.128±0.342cd
3 qRLW3 16 3.260±0.304bcd
4 qRLW5 26 3.168±0.333cd
5 qRLW2+qRLW3 4 3.649±0.167a
6 qRLW2+qRLW5 3 3.299±0.159abc
7 qRLW3+qRLW5 5 3.612±0.471ab
8 qRLW2+qRLW3+qRLW5 3 3.582±0.512ab
Table 1 Correlation coefficients among the GL, GW, GT, and RLW in the F2 population.

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

**Indicate significant at p=0.01 level, respectively.

Table 2 QTL for grain size related traits detected in the F2 population derived from a cross between Hopum and Basmati 370.

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

yLOD: The logarithm of the odds.

xPVE: Percentage of the phenotypic variance that was explained.

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

Table 3 QTL pyramiding effect on grain size related traits. Letters from a to d indicate significantly different values according to Duncan's multiple range test (α=0.05).

Means with same letters are not significantly different at p<0.05 (ANOVA followed by DMRT).

zTraits: GL; grain length, GW; grain width, GT; grain thickness, RLW; ratio of length to width.

Table 1

Table 2

Table 3

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