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RIL집단을 활용한 콩 종실의 취반특성 검정 및 연관 QTL 탐색

허진호1,2, 김지민1, 이택림1,3, 정지영1, 신일섭1, 이주석2, 강성택1,*

QTL Analysis of Seed Cooking Quality using RIL Population in Black Soybean

Jinho Heo1,2, Ji-Min Kim1, Taeklim Lee1,3, Jiyoung Jung1, Ilseob Shin1, Ju Seok Lee2, Sungtaeg Kang1,*
Korean Journal of Breeding Science 2022;54(1):25-33.
Published online: March 1, 2022

1충청남도 천안시 단대로 119 단국대학교 생명공학대학 식량생명공학전공

2충청북도 청주시 오창읍 한국생명공학연구원 바이오평가센터

3경기도 평택시 오성면 경기도종자관리소

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*Corresponding Author (E-mail: kangst@dankook.ac.kr, Tel: +82-41-550-3621)
• Received: January 24, 2022   • Revised: February 10, 2022   • Accepted: February 11, 2022

Copyright © 2022 by the Korean Society of Breeding Science

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  • In Korea, black soybeans are traditionally consumed after cooking with rice to supplement protein and oil which are lacking in rice. Seed cooking quality including seed traits after cooking with rice, which is important for consumers, were so far not comprehensively considered during the process of breeding. In this study, we first evaluated seed quality after cooking with rice, we tested the correlation between seed cooking quality and seed water absorption ratio, and we attempted to identify the Quantitative trait locus(QTL)/gene using two recombinant inbred line (RIL) populations, i.e., Daepung × Socheong2 and Daepung × Ilpumgeomjeong. Based on phenotype and correlation analyses, the main factors affecting the hardness of soybeans cooked with rice may differ between RIL population. In the Daepung × Socheong2 RIL population, one QTL associated with seed hardness after cooking with rice was identified on chromosome 11, and Glyma.11g049600, encoding peroxidase, is proposed as a candidate gene. In the Daepung × Ilpumgeomjeong RIL population, two QTLs associated with seed hardness after cooking with rice were identified on chromosomes 7 and 19, one QTL related to seed water absorption on chromosome 3, and Glyma.19g092600 encoding pectin methylesterase inhibitor are proposed as candidate genes. This is the first study on soybean cooking quality after cooking with rice, and the locations of four related QTLs were identified. The results will be of use for future development of high-quality black soybean varieties with high consumer preference using molecular breeding methods.
콩 [Glycine max (L.) Merr]은 우리나라를 비롯한 아시아 지역에서 약 3,000년 전부터 재배되어 왔으며, 특히 우리나라에서는 단백질의 주된 공급원으로서 다양한 형태로 섭취하여 왔다. 황색콩은 두부, 장류(된장, 고추장), 콩기름, 두유제품 등의 식품제조 및 가공원료로 주로 사용되고, 검정콩인 흑태와 서리태는 주로 밥과 함께 혼식 또는 콩자반으로 이용되며, 쥐눈이콩으로 불리는 서목태는 가장 작은 크기에 윤기가 나며 해독작용이 뛰어나다고 알려져 한방소재로 많이 사용되었다(Kwon 2000, Oh et al. 1992). 이러한 검정콩에는 노화예방에 효능이 있는 항산화물질인 안토시아닌, 사포닌, 비타민 E, 카로티노이드 등이 풍부하다고 알려져 있고 관련 연구도 지속적으로 수행되고있다. Takahashi et al. (2005)등은 검정콩이 황색콩보다 종피의 페놀화합물의 양이 높아 Low-Density Lipoprotein Cholesterol (LDL)작용을 억제하는 것으로 보고하였고, Lee et al. (2014)등은 검정콩의 여러 페놀물질 중 안토시아닌이 항암 및 항산화효과, 혈중 중성지방과 콜레스테롤의 저하 등 여러 생리활성 효과를 보여주었다고 보고하였다(Takahashi et al. 2005, Lee et al. 2014). 인체 암세포를 이용한 실험에서도 메탄올 추출물에 의한 암세포 증식 억제 효과를 검토한 결과, 황색대두보다는 검정콩에 의한 효과가 높았으며 흑태, 서목태, 서리태 등 검정콩 메탄올 추출물은 암세포 증식을 60~70%가량 억제효과를 보여주는 등 다양한 생리활성효과를 보인 바 있고(Lim 2010), 이외에도 검정콩가루 혼합정도가 총 폴리페놀 및 플라보노이드 함량, ABTS radical 소거활성과 DPPH radical 소거활성과 높은 정의 상관을 갖는다는 결과가 보고하였다(Woo et al. 2018).
최근에는 건강에 대한 관심이 증가하면서 곡류와 두류를 혼합한 잡곡밥의 섭취량이 늘어나면서 검정콩의 소비비중이 점차 증가하고 있는데, 식품소비행태 보고서에 따르면 2018년 잡곡 구입 항목들 중 검정콩(서리태)이 22.9%로 가장 많이 구입되었으며, 2018년에는 21.4%로 현미찹쌀(24.2%) 다음으로 많이 구입하였다는 보고가 있다(KREI 2019). 밥밑용 검정콩의 식미와 이화학적 특성에 관한 연구에서 잡곡과 두류의 영양성분을 분석한 결과 조단백질과 조지방함량, 철분과 칼슘함량이 서리태에서 가장 높았고 취반 후 경도는 가장 낮았다(Lim et al. 2003). 밥밑용 검정콩을 15시간 침지하고 1분 30초간 끓인 콩을 이용하여 밥밑용 검정콩의 관능검사와 이화학적 특성 간에 상관관계를 분석한 결과 종합적기호도는 종자의 크기와는 정의상관을 조단백 및 조섬유 함량과는 부의 상관을 보였고, 단맛, 비린 맛, 고소한 맛 등 식미는 조섬유함량 과 부의 상관을 보였다(Seo et al. 2015). 이외에도 잡곡밥에 대한 기호도 조사(Jang et al. 2012), 콩의 종류 및 조리방법에 따른 아미노산 및 단백질함량 등의 연구(Im et al. 2016), 콩의 자엽 및 종피의 성분이나 균열의 유무, 수분흡수율 등과 취반특성 간의 관계 등에 관한 연구들도 수행되어 왔다(Ma et al. 2004, Mullin & Xu 2001).
그러나 취반 형태별 품질특성에 대한 성분비교, 기호도 조사, 관능평가 등 이화학적 및 물리적 특성 연구에 비해 분자유전학적 접근과 같은 유전 및 육종 연구는 전무하다. 최근 분자마커를 이용한 유전자지도 작성 및 QTL (Quantitative Traits Loci)탐색이 여러 작물을 대상으로 진행되고 있고(Fang et al. 2016, Sandhu et al. 2018), 미국에서는 콩 품종 ‘Williams82’의 유전체 염기서열이 2010년에 해독완료 되어 콩의 유전체 정보를 제공하고 있다(Schmutz et al. 2010). 국내연구팀은 표준유전체를 참고하여 작성된 콩의 실제물리지도상에 존재하는 SNP (Single Nucleotide Polymorphism)마커들을 활용 하여 AxiomⓇ 180K Soya SNP array를 개발하였으며, 이를 통해 고밀도 유전자지도 작성 및 QTL탐색이 가능하게 된 상황에서(Lee et al. 2015) 본 연구는 검정콩의 취반특성에 연관된 유전자 탐색을 통해, 기능성 및 수량성이 우수할 뿐만 아니라 취반특성에 대한 소비자들의 요구를 충족시킬 수 있는 고품질의 밥밑콩을 육종할 수 있는 기초 자료를 제공 하고자 수행하였다.
이를 위해 본 연구에서는 (1) 종실의 취반특성 및 수분흡수율에 대한 표현형을 조사하고, 특성 간의 상관분석을 하였으며, (2) 두 RIL집단에 대한 표현형정보와 AxiomⓇ 180K Soya SNP array를 이용하여 얻은 유전형정보를 통해 유전자지도를 작성한 후 (3) 취반특성과 수분흡수율에 연관된 QTL 및 후보유전자를 탐색하였다.
실험재료
본 연구에는 내도복성이며 내병충성 특성을 가진 다수성 품종인 ‘대풍’(Park et al. 2005)을 공통 모본으로 소립 검정콩이며, 경도가 낮은 ‘소청2호’(Baek et al. 2013)을 부본으로 한 대풍×소청2호 RIL집단 107계통과 대립이며 밥밑용 콩 특성이 우수한 ‘일품검정콩’(Shin et al. 1998)을 부본으로 한 대풍×일품검정 RIL집단 103계통을 사용하였다. 두 RIL집단은 단국대학교에서 인공교배를 통한 F1을 양성 후 F2세대부터 SSD (single seed descent)방법으로 F5~F7 까지 세대를 진전시킨 후 Bulk로 세대진전한 F10 이후 세대 종자를 사용하였다.
종실 취반특성 취사 조건 및 경도, 당도 측정
종실의 취반특성 조사를 위해 압력밥솥에 쌀과 함께 취사하는 방법을 사용하였는데, 먼저 쌀 300 g을 3~4회 세척 후 채반에 걸러 5분간 물기를 제거를 하고, 4시간 침지한 30립의 종자 중 경실종자 및 피해립을 제거하고 균일한 상태의 콩 13~15립을 준비한 쌀과 함께 물 300 mL을 추가하여 압력밥솥(CJS-FC1001F, (주)쿠첸, KOREA)에 치상하였다. 1회 취반시 7품종을 임의로 배치하였고 압력밥솥의 취사방법은 ‘잡곡취사’로 선택하였다. (Figs. 1A, 1B, 1C).
취반후 종실 경도측정은 취반한 콩 13~15립 중 균일한 상태의 콩을 10립을 선발하여 과일경도계(FHR-1, NOW, JAPAN)를 이용하여 측정하였다. 취사가 끝난 직후 밥솥 내의 증기 배출 뒤 종자를 꺼낸 후 쌀과 콩이 접촉한 부위를 수직으로 경도 측정 후 평균값을 이용하였다(Fig. 1D). 측정한 값은 경도단위 환산식을 이용해 N으로 환산하였다(Park 2012).
(1)
kg⋅f kilogram-force×9.807=N
당도는 취반한 콩을 증류수로 1/10로 희석하여 믹서로 마쇄 후 1.5 mL tube에 담아 13,000 rpm, 5분 조건에서 원심분리 후 상층액 300 μL를 채취하여 굴절당도계(PAL-1 Digital Hand Held Pocket Refractometer, ATAGO, USA)를 이용하여 3반복으로 측정하였다(Fig. 1E). 측정값은 10배로 환산 후 평균값을 이용하였다.
종실 수분흡수율
종실 수분흡수율 측정은 종자를 Dry Oven (NB-901M, N- BIOTEK, KOREA)에 60~70℃ 온도에서 24시간 건조 후 30립의 무게를 측정하고, 측정한 종자를 침수 4시간 침수 후 종자 표면수를 제거한 다음 무게를 측정하여 침지 전과 후의 무게 차를 종자의 무게로 나누어 수분흡수율을 계산하였다. 실험은 3반복 실시하여 평균값을 이용하였다(Wani et al. 2017).
DNA추출 및 Genotyping
종자 파종 후 V1 시기에 식물체의 삼복엽을 채취 후CTAB (Cetyltrimethyl ammonium bromide)방법을 이용하여 genomic DNA를 추출하고(Porebski et al. 1997), 추출한 genomic DNA를 180,961개의 SNP로 구성된 Axiom® 180K Soya SNP array (Affymetrix, CA)를 이용하여 대풍, 소청2호, 일품검정콩과 대풍×소청2호 RIL집단 107계통 및 대풍×일품검정 RIL집단 103계통에 대하여 genotyping실시하였다(Lee et al. 2015).
유전자지도작성 및 QTL 분석
169,028개의 SNP 마커들 중 모부본 사이에 다형성을 보이는 SNP마커를 선발하고, QTL IciMapping V4를 Binning : Segregation Distortion 0.01, Grouping : LOD 3.0이상, Rippling : Sum of adjacent recombination fractions (SARF), Ordering : nnTwoOpt 조건으로 설정하여 유전자지도를 작성하였다(Meng et al. 2015). 작성된 유전자지도와 표현형 데이터를 활용하여 ICIM (Inclusive Composite Interval Mapping)의 방법으로 QTL분석을 실시하였다(Meng et al. 2015). 탐색한 QTL을 바탕으로 물리적 거리를 Soybase (http://soybase.org/gb2/gbrowse/, 2021)에서 확인하여 후보 유전자를 탐색하였다(Grant et al. 2010).
취반 후 종실 취반특성
두 RIL집단에서 취반특성인 종실의 취반경도(N)는 대풍×소청2호 RIL집단의 모본인 대풍은 3.46 N, 부본인 소청2호는 2.56 N을 보였고, RIL계통은 2.08~3.54 N 범위에 평균값은 2.70 N이고, 2.50~2.76 N범위에서 가장 많은 26계통이 분포하였다(Table 1, Fig. 2A). 대풍×일품검정콩 RIL집단에선 모본인 대풍은 3.46 N, 부본인 일품검정은 2.43 N을 보였으며 RIL계통은 1.89~3.73 N범위에 평균값은 2.73 N이며, 2.60~2.80 N범위에서 가장 많은 29계통이 분포하였다(Table 1, Fig. 2A). 두 RIL집단을 비교하였을 때 평균값은 유사하였고 정규분포 하였다(Fig. 2A). 따라서 취반 후 종실 경도와 관련된 유전자는 양적형질로 2개이상의 유전자가 동시에 관여할 것으로 추정되며 양집단에서 모부본을 능가하는 초월분리현상도 관찰되었다. 두 RIL집단에서 취반특성인 취반후 종실의 당도(Brix)에 대한 분포는 공통 모본인 대풍은 14.3 Brix, 소청2호는 11.0 Brix, 일품검정콩은 12.3 Brix 로 나타났고, 집단별로는 대풍×소청2호 집단은 9.0~14.3 Brix 범위에서 평균 10.9 Brix이며 10.0~11.0 Brix 범위에서 가장 많은 50계통이 분포하였다(Table 1, Fig. 2B). 대풍×일품검정콩 집단은 9.8~15.0 Brix 범위에 평균 12.3 Brix 이고, 11.0~12.0 Brix범위에서 가장 많은 36계통이 분포하였다(Table 1, Fig. 2B). 취반 후 종실의 당도(Brix)분포는 두 집단에서 모두 정규분포양상을 보였고, 범위도 유사하였으며 두 집단 모두 초월분리현상을 보였다.(Fig. 2B). 두 RIL집단에서 종실의 수분흡수율(%)에 대한 분포도를 보았을 때 모본인 대풍은 61.8%, 소청2호는 86.9%, 일품검정콩은 86.1%를 보였다. 계통별 분포를 살펴보면 대풍× 소청2호 집단은 62.4~103.7% 범위에서 평균값은 87.4%를 나타냈고, 86~89%범위에서 가장 많은 계통이 분포하였다(Table 1, Fig. 2C). 대풍×일품검정콩 집단에선 56.0~97.0% 범위에서 평균값은 79.8%를 보였고, 78.0~84.0 범위에서 가장 많은 16계통으로 분포하였다(Table 1, Fig. 2C). 두 집단 모두 정규분포 하였고 초월분리 현상도 관찰되었다.(Fig. 2C).
종실 수분흡수율 및 취반특성 간 상관분석
RIL집단에서 종실특성 간에 상관분석 결과를 보면 대풍× 소청2호 경도와 수분흡수율, 당도와 수분흡수율 간에는 높은 부의 상관, 경도와 당도는 낮은 정의 상관을 보였다. 특히, 경도와 수분흡수율 간에는 -0.697*** 가장 높은 부의 상관을, 다음으로 당도와 수분흡수율 -0.508*** 부의 상관을 보였다(Table 2). 대풍×일품검정콩 집단에서는 경도와 당도는 낮은 정의 상관, 경도와 수분흡수율, 당도와 수분흡수율 간에 모두 0.5미만으로 낮은 상관을 나타냈다(Table 2). 두 집단간에 상관분석 결과에서 집단간에 형질별 상관관계 정도가 다르게 나타나고 있어 집단별로 관여 요인의 차이가 일을 것으로 보여진다(Table 2).
종실 취반특성 및 수분흡수율 연관 QTL분석

1. 유전자지도작성

유전자지도 작성결과를 보면 대풍×소청2호 RIL집단은 지도에 표기된 marker는 10,720개로 염색체당 최소 192개, 최대 943개가 이용되었으며, 평균 536개의 marker가 활용되었다(Fig. 3A). 작성된 유전자지도는 총 2,048 cM이며 염색체당 평균 지도의 길이는 102.43 cM으로 marker간 평균거리는 0.19 cM이다(Fig. 3A). 대풍×일품검정콩 집단에서 유전자 지도 작성에 사용된 SNP marker는 9,916개로 평균 496개의 marker가 활용되었다(Fig. 3B). 작성된 유전자지도는 총 2,039 cM이며 염색체당 평균 지도의 길이는 101.98 cM으로 전체지도 내에 작성된 marker간 평균거리는 0.21 cM이다(Fig. 3B).

2. 연관 QTL 및 후보 유전자 탐색

종실 특성과 관련된 QTL탐색 결과를 보면 취반 후 경도와 관련된 QTL이 대풍×소청2호 집단은 11번염색체에서 1개의 QTL이 탐색되었고, 대풍×일품검정콩 집단은 7번, 19번 염색체에서 각각 1개의 QTL이 탐색되었다. 또한 수분흡수율과 관련된 QTL은 대풍×일품검정콩 집단의 3번 염색체에서 1개의 QTL이 탐색되었으나 전반적으로 두 집단에서 탐색된 QTL의 PVE (%)가 20미만으로 모두 다소 낮게 탐색되었다(Table 3).
QTL탐색 결과를 좀 더 자세히 살펴보면, 대풍×소청2호 집단에서 11번 염색체에서 탐색된 QTL은 AX-90402610~AX- 90476968의 두 flanking marker사이에서 탐색되었고, 물리적 위치는 3,721,311~3,999,032 bp에 위치하였으며, 그 간격은 278 kbp였다(Table 3, Figs. 4A, 4B). Soybase (http://soybase.org/gb2/gbrowse/, 2021)에서 후보 유전자를 탐색한 결과 QTL영역내에서 27개의 후보유전자가 탐색되었고, 이중 경도와 관련이 높을 것으로 추정되는 Peroxidase 51-like인 Glyma.11g049600이 탐색되었다(Table 3, Fig. 4C). 종자생합성 과정에 관여하는 것으로 알려진 Peroxidase는 콩의 종피와 상관관계가 높으며, 콩의 종피 내에 Peroxidase 활성이 높게 나타난다는 보고가 있으며(Gijzen et al. 1993), 종피와 수분흡수율간에 높은 상관관계를 나타낸 연구도 보고되었다(Mullin & Xu 2001).
대풍×일품검정콩 집단에서는 경도와 연관된 QTL이 7번, 19번 염색체에서 탐색되었고, 7번 염색체에서 탐색된 QTL은 AX-90450539~AX-90519477의 두 flacking marker사이에서 탐색되었고, 물리적 위치는 35,730,875~36,675,391 bp에 위치하며, 그 간격은 944 kbp였다(Table 3, Figs. 5A, 5B). 19번 염색체에서 탐색된 QTL은 AX-90322246~AX-90434088의 두 flanking marker사이에서 탐색되었으며, 실제 염색체 상에서는 32,353,983~ 37,152,969 bp에 위치하였고, 그 간격은 4.8 Mb였다(Table 3, Figs. 5A, 5C). Soybase (http://soybase.org/gb2/gbrowse/, 2021)에서 탐색된 영역내에 후보 유전자를 탐색한 결과 7번 염색체에서는 81개 후보유전자가 탐색되었지만 이 영역 내에서 경도와 직접적으로 연관된 후보유전자를 탐색하지 못하였다(Table 3, Fig. 5D).
19번 염색체에서는 219개의 후보유전자가 탐색되었고, 이중 WRKY53, GEM-like protein, Phosphoribosylglycinamide 등 25개의 유전자가 알려져 있다(Table 3, Fig. 5E). 후보유전자 영역에서 경도와 관련된 유전자 Pectin Methylesterase (PME)의 inhibitor인 Glyma.19g092600이 탐색되었다. PME는 취반후 콩의 자엽의 경도를 결정하는데 연관있는 유전자로 Glyma.03g028900이 보고된 바 있다(Hirata et al. 2014, Toda et al. 2015, Toda et al. 2020). 보고에 의하면 PME는 Ca2+이온이 pectin 단백질과 결합하여 세포벽의 경도를 결정하는데 영향을 준다는 연구결과가 있어(Catoire et al. 1998) 본 연구에서 발견한 후보유전자인 PME의 기능을 저해하는 유전자로 자엽의 경도와 연관 있는 유전자로 판단하였다.
수분흡수율과 연관된 QTL은 3번 염색체 AX-90430380~AX- 90519747의 두 flacking marker사이에서 탐색되었고, 물리적 위치는 34,432,251~36,200,178 bp에 위치하며, 그 간격은 1.7 Mb였다(Table 3, Figs. 6A, 6B). Soybase (http://soybase.org/gb2/gbrowse/, 2021)에서 확인하여 후보 유전자를 탐색한 결과 154개의 후보유전자가 탐색되었고, 17개의 유전자는 기능이 보고 되었으나 후보 유전자 영역 내에서 수분흡수율과 관련된 유전자를 탐색하지 못하였다(Table 3, Fig. 6C).
본 연구에서 사용한 두 개 집단에서의 QTL 분석 결과가 상이하였는데 이는 첫째, 실험재료에 의한 차이로, 실험에 이용된 RIL집단은 공통 모본을 가지고 서로 다른 부본으로 RIL 집단의 유전적배경 다르기에 실험결과의 차이가 났을 것으로 예측한다. 둘째, 기 보고된 콩들과는 달리, 본 실험은 취반한 콩을 이용하였기에 추가로 발생하는 요인이 작용하여 취반 경도에 관련된 새로운 QTL을 탐색한 것으로 사료된다. 두 RIL집단 모두 종실의 취반 당도와 관련된 QTL은 탐색되지 않았는데 이는 교배 모부본간에 당도값의 변이가 적고, 유전적인 차이가 크지 않아서 QTL탐색이 제한적이었다고 추측한다. 탐색을 통해 확인된 후보유전자들은 추후 연구에서 발현양상을 비교하고 각 특성연관 유전자임을 확인할 필요가 있다.
밥밑용 콩으로 주로 소비되는 검정콩은 지속적으로 품종개발이 되고 있으나 다수의 밥밑용 콩 품종의 육종목표는 기능성 및 수량성 향상에 초점이 맞춰져 있다. 또한, 취반 전 종실의 당도와 경도에 관한 연구는 많이 보고되어 있지만, 소비자들이 구입할 때 고려하는 취반 후 품질에 대한 연구는 미비하며 육종단계에서 충분히 반영되지 않고 있다.
본 연구는 검정콩의 취반특성에 연관된 유전자를 탐색하고자 두 RIL집단을 대상으로 취반 후 특성(경도, 당도)및 수분흡수율에 대한 표현형 조사를 하였다. 이를 바탕으로 특성간의 상관관계를 파악하고, 나아가 유전형 정보를 토대로 특성연관 QTL을 탐색하였다. 이를 요약하면 다음과 같다.

  1. 대풍×소청2호 RIL집단에서는 취반 후 경도는 2.08~3.54 N, 취반 후 당도는 9~12.67 brix, 수분흡수율은 63.62~103.86% 범위에, 대풍×일품검정 RIL집단에서는 취반 후 경도는 1.89~3.73 N, 취반 후 당도는 9.78~15.0 brix, 수분흡수율은 56.08~96.93%에 범위를 보였다. 두 집단 모두 각 특성들에 따른 분포가 정규분포를 보였다.

  2. 대풍×소청2호 RIL집단에서는 경도와 수분흡수율 간에는 -0.697***로 부의 상관, 당도와 수분흡수율간에 -0.508***로 부의 상관을 보였다. 대풍×일품검정RIL집단에서는 경도와 당도는 낮은 정의 상관, 경도와 수분흡수율, 당도와 수분흡수율 간에 각각 낮은 부의 상관을 보였으나 형질간 상관계수는 모두 0.5미만으로 낮은 상관을 나타냈다.

  3. 두 RIL집단에 대한 유전자 지도 작성 후 종실 특성과 관련된 QTL탐색결과, 취반후 경도와 관련된 QTL이 대풍×소청2호 RIL집단은 11번 염색체에서 1개, 대풍×일품검정 RIL집단은 7번, 19번 염색체에서 각각 1개씩 탐색되었고 해당 영역에서 후보 유전자를 탐색하고 이중 Peroxidase 51-like인 Glyma.11g049600, Pectin Methylesterase (PME)의 inhibitor인 Glyma.19g092600 유전자가 가장 유력한 후보 유전자로 추정되었다.

  4. 수분흡수율과 관련된 QTL의 경우, 대풍×일품검정 RIL집단은 3번 염색체에서 탐색되었고, 대풍×소청2호 RIL집단에선 탐색되지 않았다. 탐색된 영역 내에서 154개의 후보 유전자 중 강력히 유망한 유전자를 특정하지 못하였다.

본 연구 결과 검정콩의 취반 특성과 연관된 QTL 탐색과 더불어 후보유전자를 탐색하였다. 비록 대부분 PVE (%)가 20 미만으로 낮고 minor QTL이지만 본연구결과는 신품종 육성을 함에 있어 소비자들의 기호에 맞는 우수한 품질의 밥밑용 검정콩을 육종하기위한 기초자료로 사용할 수 있을 것이다.
본 연구는 농촌진흥청 바이오그린연계농생명혁신기술개발사업(과제번호:PJ015742)의 연구지원을 받아 이루어진 것으로 이에 감사드립니다.
Fig. 1
Procedure and device of measurement of seed quality after cooking with rice (A) Arrangement of materials before cooking with rice (B) Electric rice cooker (C) Arrangement of materials after cooking with rice (D) Hardness tester (FHR-1, NOW, JAPAN) (E) Digital refractometers (PAL-1, ATAGO, U.S.A).
kjbs-54-1-25-f1.jpg
Fig. 2
Distribution of seed hardness, brix after cooking with rice and water absorption in two RIL population. Daepung (P1) is represented to dashed box, Socheong2 (P2) is represented to black box, Ilpumgeomjeong (P3) is represented to white box. A : seed hardeness after cooking with rice in Daepung×Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white), B : seed brix after cooking with rice in Daepung× Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white), C : water absorption in Daepung×Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white).
kjbs-54-1-25-f2.jpg
Fig. 3
Genetic map for the two RIL populations A: Daepung×Socheong2 RIL population, B : Daepung×Ilpumgeomjeong RIL population. Black bars mean SNP marker position on each chromosome.
kjbs-54-1-25-f3.jpg
Fig. 4
Identification of QTLs and candidate gene related to seed hardness after cooking with rice in Daepung×Socheong2 RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 11, C : Candidate genes on chromosome 11.
kjbs-54-1-25-f4.jpg
Fig. 5
Identification of QTLs and candidate gene related to seed hardness after cooking with rice in Daepung×Ilpumgeomjeong RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 7, C : LOD graph on chromosome 19, D : Candidate genes on chromosome 7, E : Candidate genes on chromosome 19.
kjbs-54-1-25-f5.jpg
Fig. 6
Identification of QTLs and candidate gene related to water absorption in Daepung×Ilpumgeomjeong RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 7, C : LOD graph on chromosome 19, D : Candidate genes on chromosome 7, E : Candidate genes on chromosome 19.
kjbs-54-1-25-f6.jpg
Table 1
Summary of result for three seed quality traits after cooking with rice in two RIL populations
Table 1
Population Trait Parents RIL population
Daepung Founder Mean±SD Range
Daepung
×
Socheong2		 Cooked seed hardness (N) 3.5±0.2 2.6±0.1 2.7±0.4 2.1~3.5
Sucrose content (Brix%) 14.3±1.2 11.0±0.0 10.9±1.3 9.0~12.7
Water absorption (%) 61.8±4.3 86.9±3.3 86.7±12.0 63.6~103.9
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness (N) 3.5±0.2 2.4±0.7 2.7±0.4 1.9~3.7
Sucrose content (Brix%) 14.3±1.2 12.3±1.2 12.3±1.0 9.8~15.0
Water absorption (%) 61.8±4.3 810.0±9.8 79.9±8.3 56.1~96.9
Table 2
Correlation coefficients between three seed quality traits after cooking with rice in two RIL populations
Table 2
Population Trait Cooked seed hardness (N) Sucrose content (Brix%) Water absorption (%)
Daepung
×
Socheong2		 Cooked seed hardness (N) 1
Sucrose content (Brix%) 0.224* 1
Water absorption (%) -0.697*** -0.508*** 1
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness (N) 1
Sucrose content (Brix%) 0.450*** 1
Water absorption (%) -0.228* -0.327*** 1

Ns: not significant, p<0.05 : *, p<0.01 : **, p<0.001 : ***

Table 3
List of QTLs associated with cooked seed hardness and water absorption in two RIL populations
Table 3
RIL population Trait Chrz Markers physical
position (bp)y 		Intervalx LODw PVE (%)v Addu Number of candidate gene Published gene Potential candidate gene
Daepung
×
Socheong2		 Cooked seed hardness 11 AX-90402610
~
AX-90476968		 3,721,311
~
3,999,032		 277,721 3.3 13.6 0.1 27 3 Peroxidase
Glyma.11g049600
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness 7 AX-90450539
~
AX-90519477		 35,730,875
~
36,675,391		 944,516 3.4 6.2 -0.1 81 1 -
19 AX-90322246
~
AX-90434088		 32,353,983
~
37,152,969		 4,798,986 4.2 7.9 0.1 219 25 Pectin Methylesterase
Inhibitor
Glyma.19g092600
Water
absorption		 3 AX-90430380
~
AX-90519747		 34,432,251
~
36,200,178		 1,767,927 3.7 20.0 -2.8 154 17 -

zchromosome

yinformation from Glyma.Wm82.a2.v1 genome browser (Grant et al. 2010)

xphysical distance between two markers

wlogarithm of odds score, threshold was 2.79 from permutation test

vphenotypic variation explained by the marker

umean of the distribution (P2-P1)/2

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QTL Analysis of Seed Cooking Quality using RIL Population in Black Soybean
Korean. J. Breed. Sci.. 2022;54(1):25-33.   Published online March 1, 2022
DOI: https://doi.org/10.9787/KJBS.2022.54.1.25
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QTL Analysis of Seed Cooking Quality using RIL Population in Black Soybean
Korean. J. Breed. Sci.. 2022;54(1):25-33.   Published online March 1, 2022
DOI: https://doi.org/10.9787/KJBS.2022.54.1.25
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QTL Analysis of Seed Cooking Quality using RIL Population in Black Soybean
Image Image Image Image Image Image
Fig. 1 Procedure and device of measurement of seed quality after cooking with rice (A) Arrangement of materials before cooking with rice (B) Electric rice cooker (C) Arrangement of materials after cooking with rice (D) Hardness tester (FHR-1, NOW, JAPAN) (E) Digital refractometers (PAL-1, ATAGO, U.S.A).
Fig. 2 Distribution of seed hardness, brix after cooking with rice and water absorption in two RIL population. Daepung (P1) is represented to dashed box, Socheong2 (P2) is represented to black box, Ilpumgeomjeong (P3) is represented to white box. A : seed hardeness after cooking with rice in Daepung×Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white), B : seed brix after cooking with rice in Daepung× Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white), C : water absorption in Daepung×Socheong2 population (black), Daepung×Ilpumgeomjeong RIL population (white).
Fig. 3 Genetic map for the two RIL populations A: Daepung×Socheong2 RIL population, B : Daepung×Ilpumgeomjeong RIL population. Black bars mean SNP marker position on each chromosome.
Fig. 4 Identification of QTLs and candidate gene related to seed hardness after cooking with rice in Daepung×Socheong2 RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 11, C : Candidate genes on chromosome 11.
Fig. 5 Identification of QTLs and candidate gene related to seed hardness after cooking with rice in Daepung×Ilpumgeomjeong RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 7, C : LOD graph on chromosome 19, D : Candidate genes on chromosome 7, E : Candidate genes on chromosome 19.
Fig. 6 Identification of QTLs and candidate gene related to water absorption in Daepung×Ilpumgeomjeong RIL population. A : LOD graph on whole genome, B : LOD graph on chromosome 7, C : LOD graph on chromosome 19, D : Candidate genes on chromosome 7, E : Candidate genes on chromosome 19.

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Fig. 5

Fig. 6

QTL Analysis of Seed Cooking Quality using RIL Population in Black Soybean

Summary of result for three seed quality traits after cooking with rice in two RIL populations

Population Trait Parents RIL population
Daepung Founder Mean±SD Range
Daepung
×
Socheong2		 Cooked seed hardness (N) 3.5±0.2 2.6±0.1 2.7±0.4 2.1~3.5
Sucrose content (Brix%) 14.3±1.2 11.0±0.0 10.9±1.3 9.0~12.7
Water absorption (%) 61.8±4.3 86.9±3.3 86.7±12.0 63.6~103.9
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness (N) 3.5±0.2 2.4±0.7 2.7±0.4 1.9~3.7
Sucrose content (Brix%) 14.3±1.2 12.3±1.2 12.3±1.0 9.8~15.0
Water absorption (%) 61.8±4.3 810.0±9.8 79.9±8.3 56.1~96.9

Correlation coefficients between three seed quality traits after cooking with rice in two RIL populations

Population Trait Cooked seed hardness (N) Sucrose content (Brix%) Water absorption (%)
Daepung
×
Socheong2		 Cooked seed hardness (N) 1
Sucrose content (Brix%) 0.224* 1
Water absorption (%) -0.697*** -0.508*** 1
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness (N) 1
Sucrose content (Brix%) 0.450*** 1
Water absorption (%) -0.228* -0.327*** 1

List of QTLs associated with cooked seed hardness and water absorption in two RIL populations

RIL population Trait Chrz Markers physical
position (bp)y		 Intervalx LODw PVE (%)v Addu Number of candidate gene Published gene Potential candidate gene
Daepung
×
Socheong2		 Cooked seed hardness 11 AX-90402610
~
AX-90476968		 3,721,311
~
3,999,032		 277,721 3.3 13.6 0.1 27 3 Peroxidase
Glyma.11g049600
Daepung
×
Ilpumgeomjeong		 Cooked seed hardness 7 AX-90450539
~
AX-90519477		 35,730,875
~
36,675,391		 944,516 3.4 6.2 -0.1 81 1 -
19 AX-90322246
~
AX-90434088		 32,353,983
~
37,152,969		 4,798,986 4.2 7.9 0.1 219 25 Pectin Methylesterase
Inhibitor
Glyma.19g092600
Water
absorption		 3 AX-90430380
~
AX-90519747		 34,432,251
~
36,200,178		 1,767,927 3.7 20.0 -2.8 154 17 -
Table 1 Summary of result for three seed quality traits after cooking with rice in two RIL populations
Table 2 Correlation coefficients between three seed quality traits after cooking with rice in two RIL populations

Ns: not significant, p<0.05 : *, p<0.01 : **, p<0.001 : ***

Table 3 List of QTLs associated with cooked seed hardness and water absorption in two RIL populations

zchromosome

yinformation from Glyma.Wm82.a2.v1 genome browser (Grant et al. 2010)

xphysical distance between two markers

wlogarithm of odds score, threshold was 2.79 from permutation test

vphenotypic variation explained by the marker

umean of the distribution (P2-P1)/2

Table 1

Table 2

Table 3

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