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벼의 CAPS 마커 개발 및 줄기굵기 특성의 양적형질 유전자좌(QTL) 분석

Development of New CAPS Markers and Their Application in QTL Analysis of Stem Diameter in Rice

The Korean Journal of Breeding Science 2014;46(2):116-128.
Published online: May 31, 2014

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*Corresponding author (E-mail: thkim@rda.go.kr, Tel: +82-31-299-1641, Fax: +82-31-299-1657)
• Received: March 5, 2014   • Revised: May 12, 2014   • Accepted: May 19, 2014

© The Korean Society of Breeding Science

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  • The next generation sequencing (NGS) has been developed rapidly in recent years, paving ways of discovering vast sequence variations among germplasms. Whole-genome sequencing was performed on the genomic DNA of Milyang23 and Gihobyeo using NGS and developed new CAPS (cleaved amplified polymorphic sequence) markers based on the single nucleotide polymorphisms (SNPs) in coding sequence between these varieties. The NGS sequencing yielded sequences of 60x coverage of the Nipponbare reference genome on average. A molecular genetic map was constructed with the recombinant inbred population derived from Milyang23/Gihobyeo cross (MGRIL) integrating the newly developed 146 CAPS makers and previously reported 219 PCR-based DNA markers. This map was applied to the detection of quantitative trait loci (QTLs) for stem internode diameters, culm length and panicle length in rice with MGRIL population. A total of 4 new QTLs were detected for stem diameter traits including the first internode diameter (I1D), second internode diameter (I2D), third internode diameter (I3D), and fourth internode diameter (I4D). Among stem diameter QTLs, qI1D5 had relatively 6.09 LOD (likelihood of odds) score and explained 8.99% of total variation. Only very small portion of SNPs through re-sequencing were used in this study. Much more markers can be developed by using SNP information acquired in this study, which will enable construction of high-density genetic map and more accurate QTL analysis of important agronomical traits with MGRIL population.
작물 육종에 있어서 농업적으로 중요한 형질들은 독립적인 단일 유전자가 아니라 전체 염색체 상에 존재하는 수많은 유 전자들에 의해서 조절되는 양적형질이다. 따라서 오늘날은 이 러한 양적형질 유전자좌를 연구하기 위해서 고밀도 분자유전 지도 작성을 통해 유전자를 찾고, 그 기능을 밝히는 등의 접 근이 많이 이루어지고 있다(Li et al. 2011, Ookawa et al. 2010).
현재까지 고밀도 분자유전지도를 작성하기 위해 RFLP (restriction fragment-length polymorphism), RAPD (randomly amplified polymorphism) 및 AFLP (amplified fragment length polymorphism) 등 다양한 DNA 마커가 개발되었다. 벼에서는 자포니카 품종인 Nipponbare와 인디카 품종인 Kasalath 교배 후대 F2집단을 활용하여 2,275개 RFLP 마커들로 구성된 고 밀도 분자유전지도를 작성한 바가 있고(Harushima et al. 1998), 이는 벼 유전체 해독 프로젝트(International Rice Genome Sequencing Project, 2005)에 활용되기도 하였다.
기존의 DNA 마커들은 식별방법이 간편하지 못하고, 고비 용과 많은 소요시간이 걸린다. 게다가 출현빈도가 낮다는 한 계를 가지고 있다. 최근에 개발된 SSR (simple sequence repeat) 과 SNP (single nucleotide phymorphism)는 품종 또는 개체 간에 훨씬 높은 다형성을 보여주며, 골고루 분포하기 때문에 이용가치가 높아 표지인자로 많은 연구(McCouch et al. 1997, McCouch et al. 2002, Nasu et al. 2002, Panaud et al. 1996) 가 이루어지고 있다. 하지만 SSR은 PCR 산물의 크기가 작아 최종 판별 시 복잡하고 어려우며, 유전자칩(microarray)을 이 용한 SNP은 다량의 시료를 빠르게 분석하는데 장점을 가지 고 있지만 개발에 많은 시간과 노력이 필요하다.
최근 차세대 염기서열 분석기술(Next Generation Sequencing) 이 발달하면서 대용량의 염기서열 분석 비용이 감소하고, 속 도가 가속화되었다. 또한, 벼 유전체 해독이 완료됨으로써 표 준유전체(reference sequence)를 기반으로 전장 유전체 재염 기서열 분석(whole genome re-sequencing)을 수행하여 SNP, 삽입(insertion) 그리고 결실(deletion) 등 다양한 마커들을 대 량으로 단기간 안에 개발할 수 있게 되었다(Huang et al. 2009, Li et al. 2011). 중국에서는 재래종 벼에 대한 재염기서열 분 석을 통해 대량의 SNP를 탐색하여 고밀도의 분자유전지도를 작성하였고, 14개의 주요 농업형질에 대한 관련 유전자를 보 고한 바 있다(Huang et al. 2010).
밀양23호/기호벼의 재조합자식 유전집단(MGRIL)에 대해 RFLP, AFLP 및 SSR 마커로 구성된 유전지도가 작성되었고 (Cho et al. 1998a, Cho et al. 1998b), 최근에는 PCR을 기반으 로 분석이 용이한 STS (sequence taqqed site), InDel (Insertion-Deletion), SSR로 구성된 분자유전지도가 보고되었다(Ji et al. 2012). 또한 이 유전집단에 대해 벼의 종실, 품질 특성, 출수 일수 그리고 미립형태적 특성 등에 대한 QTL이 보고된 바가 있다(Kang et al. 1998, Kang et al. 1999, Kwak et al. 2004, Kwak et al. 2005).
벼의 다양한 주요 형질 중에 하나인 줅기굵기는 벼의 생산 성 향상과 관련되어있다. 너무 얇거나 적정 경도를 가지지 못 한 줄기는 쓰러짐에 약하고, 쓰러진 줄기는 동화작용의 장애 와 양분이동이 억제되어 수량이 감소하거나 쌀의 질이 떨어지 게 된다(Kashiwagi & Ishimaru 2004, Kashiwagi et al. 2008, Ookawa et al. 2010). 반단간 유전자로 알려진 sd-1 (Cho et al. 1994)을 비롯해 줄기와 관련된 유전자와 QTL을 활용하여 벼의 생산성과 도복 저항성이 우수한 품종을 육성할 수 있지만 (Asano et al. 2007) 이에 대한 연구가 아직 미비한 실정이다.
본 연구는 줄기굵기 관련 QTL 연구를 수행하기 위해 차세대 대량 염기서열 분석을 통해 밀양23호/기호벼 사이의 특이적 인 SNP를 CAPS (cleaved amplified polymorphic sequences) 마커로 개발한 후, PCR 기반 마커로 구성된 MGRIL 유전지도 (Ji et al. 2012)와 통합하여 분석이 용이하면서 마커 간에 균 등한 발생 분포를 갖도록 지도를 작성하였다. 유전지도를 기 반으로 줄기굵기 형질에 관한 QTL분석과 그 이용 가능성에 대해 검토하였고, 벼의 간장과 수장에 대한 QTL분석도 동시 에 수행하였다.
유전분석 집단 및 Genomic DNA 추출
본 연구에서 사용된 유전분석 집단은 통일형인 밀양23호와 자포니카인 기호벼를 각각 모본과 부본으로 하여 교배된 162 계통의 재조합 자식계통(RILs, Recombinant Inbred Lines)으 로 구성되어있다. 이 집단은 1988년도에 교배되어 F2를 SSD (single seed descent)법으로 F6세대에 걸쳐 양성하였고, 그 후 F7세대부터는 계통당 일렬 재배하여 일계통 일종자법을 통 해 20세대 이상 세대진전하면서 형질을 고정하였다. 각 계통의 어린 잎 조직을 채취하여 Genomic DNA를 추출하였으며, 그 방법은 Ji et al. (2012)이 이미 보고한 바와 같다.
NGS 대량 염기서열 분석
밀양23호와 기호벼의 잎을 채취하여 DNeasy Plant Maxi Kit (Qiagen)를 사용하여 제공된 매뉴얼에 따라 DNA를 추출 하였으며, 추출된 DNA는 차세대 염기서열 분석에 사용하였다. 두 모 본의 벼 재염기서열 분석을 위해 Illumina Hiseq 2000 을 이용하여 paired-end sequencing을 수행하였으며, 생성된 리드의 길이는 101 bp이다. 리드는 1% 미만의 오류율을 갖는 Phred Quality Value ≥ Q20을 사용하였다. 두 모본의 염기 서열은 CLC Assembly Cell (ver. 3.3.2, http://www.clcbio.com)를 이용하여 표준유전체(Nipponbare)에 조립한 후, 변이 분석을 하였다(Jeong et al. 2013).
표준유전체는 전체 염기서열이 밝혀져 있는 Oryza sativa L. cv. Nipponbare (MSU Rice Genome Annotation Project, Pseudomolecules Build 7.0)이며, 유전체 구조를 분석하기 위한 annotation 정보는 RAP-DB(http://rapdb.dna.affrc.go.jp/) 을 이용하였다.
SNP 프라이머 제작 및 PCR 수행
차세대 염기서열 분석방법으로 해독된 양친의 유전체 염기 서열에서 CDS (coding sequence)에 존재하고, 두 품종 사이 에서 특이적으로 나타나는 SNP 중 4개(EcoRI, Hind III, Pst I, XhoI) 제한효소자리에 위치한 SNP를 후보 CAPS 마커 로 개발하였다. 이 후, 선별한 SNP를 기준으로 Vector NTI 9.0.0 프로그램을 활용하여 프라이머를 제작하였다.
PCR 반응을 위해서 반응액의 총 부피는 20 μl로 하였으며, 10X Buffer 2 μl, dNTP 10 mM 0.4 μl와 Taq polymerase 0.5 unit을 사용하였고, PCR 증폭조건은 95°C에서 5분간 denaturation 후, 95°C 30초, 58°C~62°C 30초, 72°C 1분30 초로 총 35cycle을 실행하고, 72°C 10분으로 마지막 extension 을 하였다.
CAPS 마커 개발 및 유전자형 분석
밀양23호와 기호벼의 gDNA를 주형으로 앞서 개발한 SNP 프라이머로 PCR을 수행한 뒤, 5 μl의 PCR product, 1 μl의 10X buffer, 5U의 제한효소 총 10 μl 반응액으로 37°C에서 2시간 동안 제한 효소를 처리하였다. 제한효소 처리 후 1.2% 아가로스 젤에 전기영동 한 뒤, 모본간 다형성을 검정하였다. 다형성을 보인 마커들로 MGRIL 집단의 유전분석을 행하였다.
물리 및 유전지도 작성
이 집단의 물리지도 작성을 위하여 기존에 보고된 219개의 PCR을 기반으로 한 STS, InDel, SSR 마커(Ji et al. 2012)를 개발한 CAPS 마커와 통합하였다. DNA 마커들의 프라이머 시퀀스 정보와 위치정보를 이용하여 벼 유전체상에서의 위치 를 파악하여 물리지도를 작성하였다. 이를 위해 표준유전체인 Nipponbare (build 7.0)의 위치정보를 사용하였다.
유전지도 작성을 위하여 MapDisto 프로그램(Mathias 2012) 을 이용하여 유전자형데이터를 분석하였다. MGRIL 집단의 유전형을 입력한 후, “Draw all sequence” 메뉴로 각 연관군 의 마커들간의 유전적거리를 산출하여 유전지도를 작성하였 는데, 그 때 Kosambi 함수식을 적용하였다. 최종 유전지도는 MapChart 프로그램(Voorrips 2002)을 이용하였다.
표현형 조사 및 QTL 분석
QTL 분석을 위하여 공시재료에 대한 줄기굵기, 간장(Culm length) 그리고 수장(Panicle length)을 조사하였다. 줄기굵기 형질은 1절굵기(First internode), 2절굵기(Second internode), 3절굵기(Third internode) 그리고 4절굵기(Fourth internode) 로 총 4마디를 버니어캘리퍼스를 이용하여 계통당 5개체씩 조 사하였다. 이삭목과 첫 번째 마디 사이를 1절로 정하고, 그 아 래 마디를 따라 마디 사이를 각각 2절, 3절 그리고 4절로 정하 였다. 줄기굵기는 마디 사이 중앙에서 가장 두꺼운 부분을 측 정하였다. QTL 분석은 Windows QTL Cartographer V2.5 (Wang et al. 2007)를 이용하였으며, 분석 시 composite interval mapping(CIM)을 이용하였다. 각 형질 별로 95% 유의수준으 로 1,000회의 치환을 실시하여 유의성 있는 LOD (logarithm of the odds) 값을 채택하였다.
NGS 대량 염기서열 및 mapping
재조합자식 유전집단의 모본인 밀양23호와 기호벼를 Nipponbare 서열(373Mbp)을 기준으로 재염기서열 분석을 실 시하였다. 그 결과, 밀양23호는 Nipponbare 서열대비 69.6x, 기호벼는 68.5x의 염기서열을 결정하였다(Table 1). 두 모본의 결정된 염기서열을 이용하여 assembly를 수행한 결과, 표준유 전체인 Nipponbare 서열에 mapping된 read들의 평균 depth 는 밀양23호 55.7x, 기호벼 63.3x이며, 두 모본과 Nipponbare 사이의 mapping 영역의 비율(coverage)은 각각 밀양23호 78.3%, 기호벼 90.5%를 보였다(Table 2).
Table 1
Result of whole genome re-sequencing of Milyang23 and Gihobyeo
Table 1
Variety Total Bases (bp) Depth (X) Read Count GC (%) Q20z (%) Q30y (%)

Milyang23 26,590,233,034 69.6 263,269,634 44.92 93.26 86.27
Gihobyeo 26,150,864,147 68.5 258,919,447 43.82 93.45 86.51

zA quality score of 20 represents an error rate of 1 in 100, with a corresponding call accuracy of 99%,

yA quality score of 30 represents an error rate of 1 in 1000, with a corresponding call accuracy of 99.9%.

Table 2
Result of read mapping and assembly onto Nipponbare reference genome
Table 2
Variety Mapped reads (#) Mapped reads (%) Depth (X) All mapped nucleotide (bp) Coverage (%)

Milyang23 205,997,634 78.25 55.7 344,161,476 92.05
Gihobyeo 234,289,038 90.49 63.3 365,807,427 97.84
변이 탐색 및 후보 CAPS 마커 개발
Nipponbare 서열과 두 모본 간의 변이(SNP, Insertion, Deletion)를 분석하였다. 밀양23호와 기호벼의 탐색된 변이는 각각 1,564,167개와 162,631개였으며, 이 중 두 모본 사이의 특이적으로 나타나는 변이는 밀양23호 1,237,160개, 기호벼 87,271개였다. 이러한 변이의 차이는 자포니카 품종인 기호벼 와는 다르게 자포니카와 인디카 통일형 품종인 밀양23호와 자포니카 품종인 Nipponbare의 염기서열 차이로 여겨진다. 이 변이들을 RAP-DB에서 제공하는 유전체 구조 정보에 따라 un-translated region (UTR), intron, Inter-region 및 coding region sequence (CDS)로 구분하였다. CDS에 존재하는 두 모본 간의 특이적인 SNP 중 제한효소(EcoRI, Hind III, Pst I, XhoI)자리에 위치한 SNP를 선정하여 후보 CAPS 마커 로 개발하였다. 최종적으로 146개 SNP를 분자표지 개발에 사용하였다(Fig. 1).
Fig. 1.
Example of genotyping lines in the MGRIL population. The used marker is RS0460. The PCR product size was 585bp, and Milyang23 allele was divided to 230bp and 355bp fragments after restriction enzyme digestion. ‘MY’ is Milyang23 and ‘GH’ is Gihobyeo. Each of right and left ends is size marker.
KJBS-46-116_F1.gif
유전지도 및 물리지도 작성
앞서 개발된 146개의 CAPS 마커를 이용하여 MGRIL의 유 전자형을 검정하였다. 이 집단의 유전지도 작성을 위해서 기 존에 PCR을 기반으로 개발된 219개의 DNA 마커와 통합하 였다(Ji et al. 2012)(Table 3). 146개의 CAPS 마커를 포함하 여 총 365개의 DNA 마커를 이용하여 유전지도를 작성하였 다. 총 유전거리는 1,572 cM이었으며, 마커 간의 평균 거리는 4.6 cM이었다(Fig. 3). Cho et al. (1998b)가 보고한 MGRIL 집단의 유전지도의 마커 간의 거리가 3.4 cM인 점을 비추어 볼 때 이 유전지도의 마커 밀도는 높지 못하지만, NGS로 분 석된 SNP를 기반으로 개발된 CAPS 마커는 빠른 시간에 효 율적인 분석이 가능하며, 분석이 용이한 PCR 기반 마커들로 구성되었다는 장점을 가지고 있다.
Table 3
The list of DNA markers comprising the genetic and physical map constructed in this study
Table 3
Marker type no. of markers Marker list

InDelz 36 R1M7, R1M30, R1M37, R1M47, R2M10, R2M37, R2M50, R3M10, R3M23, R3M37, R4M17, R4M43, R4M50, R5M13, R5M20, R5M30, R6M14, R6M30, R6M44, R7M7, R7M20, R7M37, R8M23, R8M33, R8M46, R9M10, R9M20, R9M30, R9M42, R10M10, R10M17, R10M30, R10M40, R11M17, R11M40, R12M43

RTMz 8 RTM4211 , RTM10147, RTM3211, RTM5697, RTM3550, RTM10742, RTM9188, RTM5516

STSz 88 STS01007, STS01009, STS01017, STS01021, STS01031, STS01039, STS01043, STS02012, STS02017, STS02025, STS02027, STS02030, STS02033, STS02036, STS02038, STS03009, STS03020, STS03025, STS03032, STS03036, STS03040, STS04001, STS04007, STS04009, STS04017, STS04024, STS04037, STS04042, STS05018, STS05020, STS05025, STS05027, STS05028, STS05037, STS05043, STS05045, STS05048, STS06013, STS06019, STS06023, STS06037, STS06040, STS07005, STS07010, STS07015, STS07021, STS07023, STS07025, STS07029, STS07036, STS07040, STS08001, STS08003, STS08005, STS08008, STS08019, STS08021, STS08023, STS08029, STS08033, STS08035, STS08038, STS08043, STS09003, STS09004, STS09007, STS09011, STS09031, STS09033, STS09035, STS09036, STS09048, STS10003, STS10005, STS10010, STS10014, STS10017, STS10037, STS11016, STS11019, STS11025, STS11039, STS12008, STS12011, STS12012, STS12019, STS12023, STS12030

SSRz 87 RM1 , RM10, RM101, RM1036, RM11, RM1155, RM1227, RM12368, RM1247, RM1300, RM1370, RM1375, RM167, RM16789, RM17, RM17303, RM17377, RM17960, RM17962, RM1880, RM19218, RM19620, RM201, RM205, RM20882, RM212, RM2136, RM214, RM224, RM226, RM22608, RM22630, RM22694, RM234, RM23736, RM23779, RM240, RM242, RM246, RM247, RM253, RM25366, RM257, RM26062, RM276, RM277, RM27970, RM28400, RM286, RM287, RM3, RM3170, RM3199, RM3252, RM332, RM3394, RM3472, RM3481, RM349,RM3664, RM3765, RM401, RM408, RM420, RM4355, RM44, RM4771, RM5055, RM5349, RM536, RM5526, RM5608, RM5633, RM5753, RM5807, RM5814, RM5907, RM6367, RM6467, RM6775, RM6840, RM6841, RM6842, RM7000, RM7389, RM7492, RM7631

SNP 146 R1S1, R1S2, R1S3, R1S4, R1S5, R1S6, R1S7, R1S8, R1S9, R1S10, R1S11, R1S12, R1S13, R1S14, R1S15, R1S16, R1S17, R1S18, R1S19, R1S20, R1S21, R1S22, R1S23, R1S24, R1S25, R2S26, R2S27, R2S28, R2S29, R2S30, R2S31, R2S32, R2S33, R2S34, R2S35, R2S36, R2S37, R2S38, R2S39, R2S40, R2S41, R2S42, R2S43, R2S44, R2S45, R2S46, R3S47, R3S48, R3S49, R3S50, R3S51, R3S52, R3S53, R3S54, R3S55, R3S56, R4S57, R4S58, R4S59, R4S60, R4S61, R4S62, R4S63, R4S64, R4S65, R4S66, R4S67, R4S68, R4S69, R4S70, R4S71, R5S72, R5S73, R5S74, R5S75, R5S76, R5S77, R5S78, R5S79, R6S80, R6S81, R6S82, R6S83, R6S84, R6S85, R6S86, R6S87, R6S88, R6S89, R6S90, R6S91, R6S92, R6S93, R7S94, R7S95, R7S96, R7S97, R7S98, R7S99, R7S100, R7S101, R7S102, R8S103, R8S104, R8S105, R8S106, R8S107, R8S108, R8S109, R8S110, R8S111, R8S112, R9S113, R9S114, R9S115, R9S116, R9S117, R10S118, R10S119, R10S120, R10S121, R10S122, R10S123, R10S124, R10S125, R11S126, R11S127, R11S128, R11S129, R11S130, R11S131, R11S132, R11S133, R11S134, R11S135, R11S136, R11S137, R12S138, R12S139, R12S140, R12S141, R12S142, R12S143, R12S144, R12S145, R12S146

zJi et al. (2012).

Fig. 3.
The genetic map constructed in this study with MGRIL population. Chromosome numbers are indicated on top of chromosome, the name of each marker is at the right side of chromosome, and genetic distance of each marker from the first marker at the top of each chromosome is at the left side. Genetic distance, measured as centimorgan or cM, is calculated using Kosambi function.
KJBS-46-116_F3.gif
유전지도 분석에 통합한 DNA 마커의 primer 서열을 표준 유전체로 사용한 nipponbare genome (build7.0)에 mapping 하여 각 마커들의 벼 유전체상에서의 물리적인 위치를 파악 한 후, 물리지도를 작성하였다. 이 물리지도에서 마커 간의 물 리적 거리의 합은 361 Mbp이었으며, 이를 총 유전거리로 나 누어 구한 1 cM당 평균 물리적거리는 230 kbp이었다. 기존 의 보고된 벼 물리지도와 유전지도의 통합분석연구 결과에서 1 cM당 244 kbp이었음을 미루어 볼 때(Chen et al. 2002), 이번에 작성된 MGRIL 유전지도와 물리지도에 근거하여 특 정한 1 cM당 평균 물리적 거리가 정확도가 높은 수치라고 할 수 있다(Fig. 2).
Fig. 2.
The physical map showing the physical location of markers used in this study. Chromosome numbers are indicated on top of chromosome, the name of each marker is at the right side of chromosome, and physical distance of each marker from the first marker at the top of each chromosome is at the left side. The unit of physical distance is mga base pairs (Mbp).
KJBS-46-116_F2.gif
MGRIL 표현형 상관관계 및 QTL 분석
밀양23호와 기호벼 그리고 MGRIL에 대해 1절굵기(I1D), 2절굵기(I2D), 3절굵기(I3D), 4절굵기(I4D), 간장(CL), 그리 고 수장(PL)의 표현형을 조사하였다(Fig. 4). 조사된 형질의. (Fig. 5).
Fig. 4.
Frequency distribution of the six traits measured in MGRILs. Mean of the parents are indicated at the top of each histogram, with each arrows representing Gihobyeo and Milyang23, respectively.
KJBS-46-116_F4.gif
Fig. 5.
Position of QTLs for the four traits measured in this study with MGRIL population. The locations of the QTLs for the first internode diameter (I1D), third internode diameter (I3D), fourth internode diameter (I4D) and culm length (CL) were shown on the genetic map.
KJBS-46-116_F5.gif
변이 분포는 연속적인 정규분포를 보였으며, 모본의 범위 를 벗어나는 초월분리현상을 나타냈다. 6가지 형질간의 상관 관계 분석한 결과, 수장과 4개의 줄기굵기에 대해 유의성 있 는 상관관계를 확인할 수 있었고, 줄기의 굵기가 굵을수록 수 장의 길이가 길어지는 경향을 나타내었다. 간장과 줄기굵기는 유의성이 낮은 상관관계를 갖거나 나타나지 않는 것을 확인 할 수 있었다(Table 4).
Table 4
Correlation coefficients among the six traits measured in the MGRIL population
Table 4
Trait I1D I2D I3D I4D CL PL

I1Dz 0.8441** 0.6546** 0.6025** 0.0557 (nsy) 0.5298**
I2D 0.8578** 0.7987** 0.1154 (ns) 0.5264**
I3D 0.9347** 0.3184** 0.4964**
I4D 0.3602** 0.4688**
CL 0.2343**

**p < 0.01

zI1D=first internode diameter; I2D=second internode diameter; I3D=third internode diameter; I4D=fourth internode diameter; CL=culm length; PL=Panicle length

yns, not significant

통합된 유전지도를 바탕으로 하여 6가지 형질에 대한 총 6 개 새로운 QTL이 탐색되었다. 그 중 2개는 간장형질 관련 QTL이며, 이들을 각각 qCL5, qCL12로 명명하였다. qCL5은 LOD 값이 3.51로 밀양23호의 allele가 간장을 2.72cm 증가 시키며, 간장 표현형 변이는 4.24%였다(Table 5). qCL12은 LOD 값이 3.09로 밀양23호의 allele가 간장을 2.47cm 증가시 키며, 간장 표현형 변이는 3.59%였다(Table 5). 이 외에도 이 전에 동일집단을 대상으로 분석된 간장형질 관련 QTL (Cho et al. 2007)이 1번과 8번 염색체에서 탐색되었다(Table 5). 특히, 1번 염색체에서 확인한 QTL은 알려져 있는 밀양23호의 반왜성 단간유전자인 sd-1에 존재했다(Ashikari et al. 2002, Cho et al. 1994, Monna et al. 2002, Sasaki et al. 2002, Spielmeyer et al. 2002)(Table 5).
Table 5
Characteristics of QTLs detected for six traits with MGRILs
Table 5
Traits No. QTL namez Chry Position LODw Additive effectv R2
Interval marker Reference
(cM)x (%)u

First Internode 1 1 27.73 12.56 0.15 21.19 RS014, RM1 sdm1 (Kashiwagi et al. 2008)

2 qI1D1 1 196.62 3.26 -0.07 5.02 RS0124, RS0125

3 qI1D5 5 84.14 6.09 0.10 8.99 RM6841, RS0578

4 6 119.17 3.75 0.08 6.06 RS0693, RM5814 Kashiwagi & Ishimaru (2004)

Second Internode 5 1 27.73 5.86 0.17 12.18 RS014, RM1 sdm1 (Kashiwagi et al. 2008)

6 6 110.87 6.10 0.18 6.31 RS0691, RM1370 Kashiwagi & Ishimaru (2004)

7 7 57.00 3.19 0.13 6.53 STS07015, RS0798 sdm7 (Kashiwagi et al. 2008)

Third Internode 8 qI3D1 1 192.10 3.17 -0.18 6.53 STS01039, RS0124

9 4 74.84 4.12 -0.20 7.76 RS0462, RS0463 qCD-4 (Wang et al. 2011)

10 7 113.39 6.45 0.26 12.94 RM1370, RS0692 Kashiwagi & Ishimaru (2004)

Fourth Internode 11 qI4D1 1 193.10 4.61 0.25 9.97 STS01039, RS0124

12 6 107.87 7.47 0.34 19.25 RS0691, RM1370 Kashiwagi & Ishimaru (2004)

13 12 20.24 3.22 0.19 5.95 RS12140, STS12019 sdm12 (Kashiwagi et al. 2008),

Culm length sd-1 (Ashikari et al. 2002),
Cho et al. 1994,
14 1 171.43 31.14 -9.99 58.23 RS0119, RS0120 Monna et al. 2002,
Sasaki et al. 2002,
(Spielmeyer et al. 2002)

15 qCL5 5 41.08 3.51 2.72 4.24 R5M13, RS0573

16 8 12.16 5.37 3.33 6.44 STS08005, STS08008 pl8 (Xiao et al. 1995),
ph8 (Xiong et al. 1999)

17 qCL12 12 94.2 3.09 2.47 3.57 RS12143, RM1300

Panicle length 18 1 134.68 3.32 0.64 6.59 R1M37, RS0115 pl1 (Xiong et al. 1999),
pl1.1 (Septiningsih et al. 2003)

19 3 41.08 5.44 0.84 11.03 R3M10, RTM5697 Pl3.1 (Septiningsih et al. 2003),
pl3b (Zhuang et al. 1997)

zQTLs were newly detected and tentatively named

yThe number of chromosome

xPosition of QTL from the top of each chromosome

wThe logarithm of the ratio of two likelihoods

vPositive and negative values indicated additive effect contributed by the alleles of Milyang23 and Gihobyeo, respectively

uPercentage of variance explained by each QTL.

수장 형질 관련 QTL에 대해 새롭게 나타난 것은 없었고, 동일집단에 대해 이전에 보고된 2개 QTL이 본 분석에서 확 인되었다(Table 5). 이는 NGS 기반 CAPS 마커로 개발된 유 전지도가 정확하며, 신규로 개발된 QTL 또한 높은 신뢰성을 갖는다는 것을 보여준다.
밀양23호/기호벼의 재조합자식 유전집단에 대한 줄기굵기 QTL 연구는 아직 보고된 바가 없으며, 본 연구를 통해서 총 4개 줄기굵기 관련 QTL이 새롭게 탐색되었다(Table 5). 이 중 1절굵기 형질 관련 QTL이 두 개 나타나 각각 qI1D1과 qI1D5 로 명명하였다. qI1D1은 1번 염색체의 RS0124-RS0125사이 에서 탐색되었고, 이 QTL의 1절굵기 표현형 변이는 5.02% 였다. 기호벼의 allele가 1절굵기를 0.07cm 증가시키고, LOD 값은 3.26이었다(Table 5). qI1D5는 5번 염색체의 RM6841-RS0578사이에서 탐색되었고, 이 QTL의 1절굵기 표현형 변 이는 8.99%였다. 밀양23호의 allele가 1절굵기를 0.10cm 증 가시키고, LOD 값은 새롭게 찾아진 QTL 중 가장 높은 값인 6.09였다(Table 5). 또한, 3절굵기 관련 형질에 대해 1개 QTL 이 확인되었고, 이를 qI3D1로 명명하였다. 이 QTL은 3번 염 색체의 STS01039-RS0124사이에서 탐색되었고, 3절굵기의 표현형 변이는 6.53%였다. 이 QTL의 LOD 값은 3.17이며, 기호벼의 allele가 3절굵기를 0.18cm 증가시켰다(Table 5). 4 절굵기 형질에 대한 1개 QTL이 탐색되었고, 이를 qI4D1로 명 명하였다. 이 QTL은 1번 염색체의 STS01039-RS0124사이에 서 탐색되었고, 4절굵기 표현형 변이는 9.97% 였다. 이 QTL 의 LOD 값은 4.61이며, 기호벼의 allele가 4절굵기를 0.25cm 증가시켰다(Table 5).
qI1D1, qI3D1, qI4D1은 1번 염색체의 유사한 지역에서 탐 색되었다. 특히, 3절과 4절굵기 관련 QTL은 동일한 마커 사 이에서 탐색되었다. 하지만 줄기굵기 측정과 QTL 분석이 독 립적으로 이루어졌으며, 여러 형질이 하나의 유전자에 의해 지 배 될 수도 있지만, 서로 다른 유전자에 의해 지배될 수도 있 기 때문에 서로 다른 QTL로 임의 명명하였다. 각각의 형질이 실제 다른 유전자에 의한 것인지 추후 연구가 요구된다. 기존 에 보고된 줄기굵기 형질 관련 QTL이 이번 결과에서도 유사 한 위치에서 나타났다. 하지만 줄기굵기 측정방법과 교배 집단 등이 다르므로 이에 대한 연구는 더 필요하다(Kashiwagi & Ishimaru 2004, Kashiwagi et al. 2008, Wang et al. 2011).
이러한 QTL 분석 결과는 유용형질들과 DNA 표지인자간 의 연관관계를 규명하여 gene tagging을 하고, 이 표지인자를 이용하여 실질적인 육종을 수행하는데 기반이 될 것으로 생 각된다.
최근 급속하게 발달한 차세대 유전체분석기술을 기반으로 밀양23호와 기호벼의 유전체 서열을 분석하고, 새로운 CAPS 마커를 개발하였다. NGS를 통해 Nipponbare 유전체 길이의 60 배수만큼 염기서열을 결정하였고, CDS 안에서 두 품종간 특이적으로 나타나는 SNP를 CAPS 마커로 이용하였다. 새롭 게 개발된 146개 CAPS 마커와 기존의 보고된 219개 마커를 통합하여 총 365개의 마커로 밀양23호/기호벼의 재조합자식 유전집단에 대해 분자 유전지도를 작성하였다. 벼의 줄기굵기 와 간장 그리고 수장에 관한 QTL을 탐색한 결과, 총 19개의 유의성이 있는 QTL을 찾을 수 있었다. 이 중에 4개 줄기굵기 형질 관련 QTL과 2개 간장 형질 관련 QTL이 기존에 보고되 지 않은 새로운 QTL이었다. 그 줄기굵기 QTL 중 가장 큰 LOD값을 갖는 qI1D5는 5번 염색체에서 탐색되었으며, 1절 굵기 표현형 변이는 8.99%였다. 또한, 간장관련 QTL 중 가 장 큰 LOD 값을 갖는 qCL5은 5번 염색체에서 탐색되었고, 이 QTL의 간장 표현형 변이는 4.24%였다. 재염기서열을 통 해 밝혀진 SNP 중 소수만이 본 연구에 사용되었다. 향후 본 연구에서 밝혀진 SNP 정보를 이용한다면 더 많은 마커를 개 발하여, 고밀도 유전지도 작성이 가능할 것이다. 더 나아가 MGRIL을 이용하여 농업적으로 유용한 형질에 대해 더 정확 한 QTL 분석과 유용유전자의 개발이 가능하게 될 것이다.
본 연구는 농촌진흥청 어젠다 1-5-15(과제번호: PJ010089 012014) “벼 고밀도 유전자지도 기반 생산성관련 분자마커 개발” 과제 및 어젠다 5-18-63(과제번호: PJ008215022014) “벼 재염기서열 및 분자마커개발을 통한 유용유전자 개발” 과제의 지원에 의해 수행되었다. 또한 본 연구는 농촌진흥청 학•연•산 협동과정 사업의 지원을 받았다.
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Development of New CAPS Markers and Their Application in QTL Analysis of Stem Diameter in Rice
Korean. J. Breed. Sci.. 2014;46(2):116-128.   Published online June 30, 2014
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Development of New CAPS Markers and Their Application in QTL Analysis of Stem Diameter in Rice
Korean. J. Breed. Sci.. 2014;46(2):116-128.   Published online June 30, 2014
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Development of New CAPS Markers and Their Application in QTL Analysis of Stem Diameter in Rice
Image Image Image Image Image
Fig. 1. Example of genotyping lines in the MGRIL population. The used marker is RS0460. The PCR product size was 585bp, and Milyang23 allele was divided to 230bp and 355bp fragments after restriction enzyme digestion. ‘MY’ is Milyang23 and ‘GH’ is Gihobyeo. Each of right and left ends is size marker.
Fig. 3. The genetic map constructed in this study with MGRIL population. Chromosome numbers are indicated on top of chromosome, the name of each marker is at the right side of chromosome, and genetic distance of each marker from the first marker at the top of each chromosome is at the left side. Genetic distance, measured as centimorgan or cM, is calculated using Kosambi function.
Fig. 2. The physical map showing the physical location of markers used in this study. Chromosome numbers are indicated on top of chromosome, the name of each marker is at the right side of chromosome, and physical distance of each marker from the first marker at the top of each chromosome is at the left side. The unit of physical distance is mga base pairs (Mbp).
Fig. 4. Frequency distribution of the six traits measured in MGRILs. Mean of the parents are indicated at the top of each histogram, with each arrows representing Gihobyeo and Milyang23, respectively.
Fig. 5. Position of QTLs for the four traits measured in this study with MGRIL population. The locations of the QTLs for the first internode diameter (I1D), third internode diameter (I3D), fourth internode diameter (I4D) and culm length (CL) were shown on the genetic map.
Development of New CAPS Markers and Their Application in QTL Analysis of Stem Diameter in Rice

Result of whole genome re-sequencing of Milyang23 and Gihobyeo

Variety Total Bases (bp) Depth (X) Read Count GC (%) Q20z (%) Q30y (%)

Milyang23 26,590,233,034 69.6 263,269,634 44.92 93.26 86.27
Gihobyeo 26,150,864,147 68.5 258,919,447 43.82 93.45 86.51

A quality score of 20 represents an error rate of 1 in 100, with a corresponding call accuracy of 99%,

A quality score of 30 represents an error rate of 1 in 1000, with a corresponding call accuracy of 99.9%.

Result of read mapping and assembly onto Nipponbare reference genome

Variety Mapped reads (#) Mapped reads (%) Depth (X) All mapped nucleotide (bp) Coverage (%)

Milyang23 205,997,634 78.25 55.7 344,161,476 92.05
Gihobyeo 234,289,038 90.49 63.3 365,807,427 97.84

The list of DNA markers comprising the genetic and physical map constructed in this study

Marker type no. of markers Marker list

InDelz 36 R1M7, R1M30, R1M37, R1M47, R2M10, R2M37, R2M50, R3M10, R3M23, R3M37, R4M17, R4M43, R4M50, R5M13, R5M20, R5M30, R6M14, R6M30, R6M44, R7M7, R7M20, R7M37, R8M23, R8M33, R8M46, R9M10, R9M20, R9M30, R9M42, R10M10, R10M17, R10M30, R10M40, R11M17, R11M40, R12M43

RTMz 8 RTM4211 , RTM10147, RTM3211, RTM5697, RTM3550, RTM10742, RTM9188, RTM5516

STSz 88 STS01007, STS01009, STS01017, STS01021, STS01031, STS01039, STS01043, STS02012, STS02017, STS02025, STS02027, STS02030, STS02033, STS02036, STS02038, STS03009, STS03020, STS03025, STS03032, STS03036, STS03040, STS04001, STS04007, STS04009, STS04017, STS04024, STS04037, STS04042, STS05018, STS05020, STS05025, STS05027, STS05028, STS05037, STS05043, STS05045, STS05048, STS06013, STS06019, STS06023, STS06037, STS06040, STS07005, STS07010, STS07015, STS07021, STS07023, STS07025, STS07029, STS07036, STS07040, STS08001, STS08003, STS08005, STS08008, STS08019, STS08021, STS08023, STS08029, STS08033, STS08035, STS08038, STS08043, STS09003, STS09004, STS09007, STS09011, STS09031, STS09033, STS09035, STS09036, STS09048, STS10003, STS10005, STS10010, STS10014, STS10017, STS10037, STS11016, STS11019, STS11025, STS11039, STS12008, STS12011, STS12012, STS12019, STS12023, STS12030

SSRz 87 RM1 , RM10, RM101, RM1036, RM11, RM1155, RM1227, RM12368, RM1247, RM1300, RM1370, RM1375, RM167, RM16789, RM17, RM17303, RM17377, RM17960, RM17962, RM1880, RM19218, RM19620, RM201, RM205, RM20882, RM212, RM2136, RM214, RM224, RM226, RM22608, RM22630, RM22694, RM234, RM23736, RM23779, RM240, RM242, RM246, RM247, RM253, RM25366, RM257, RM26062, RM276, RM277, RM27970, RM28400, RM286, RM287, RM3, RM3170, RM3199, RM3252, RM332, RM3394, RM3472, RM3481, RM349,RM3664, RM3765, RM401, RM408, RM420, RM4355, RM44, RM4771, RM5055, RM5349, RM536, RM5526, RM5608, RM5633, RM5753, RM5807, RM5814, RM5907, RM6367, RM6467, RM6775, RM6840, RM6841, RM6842, RM7000, RM7389, RM7492, RM7631

SNP 146 R1S1, R1S2, R1S3, R1S4, R1S5, R1S6, R1S7, R1S8, R1S9, R1S10, R1S11, R1S12, R1S13, R1S14, R1S15, R1S16, R1S17, R1S18, R1S19, R1S20, R1S21, R1S22, R1S23, R1S24, R1S25, R2S26, R2S27, R2S28, R2S29, R2S30, R2S31, R2S32, R2S33, R2S34, R2S35, R2S36, R2S37, R2S38, R2S39, R2S40, R2S41, R2S42, R2S43, R2S44, R2S45, R2S46, R3S47, R3S48, R3S49, R3S50, R3S51, R3S52, R3S53, R3S54, R3S55, R3S56, R4S57, R4S58, R4S59, R4S60, R4S61, R4S62, R4S63, R4S64, R4S65, R4S66, R4S67, R4S68, R4S69, R4S70, R4S71, R5S72, R5S73, R5S74, R5S75, R5S76, R5S77, R5S78, R5S79, R6S80, R6S81, R6S82, R6S83, R6S84, R6S85, R6S86, R6S87, R6S88, R6S89, R6S90, R6S91, R6S92, R6S93, R7S94, R7S95, R7S96, R7S97, R7S98, R7S99, R7S100, R7S101, R7S102, R8S103, R8S104, R8S105, R8S106, R8S107, R8S108, R8S109, R8S110, R8S111, R8S112, R9S113, R9S114, R9S115, R9S116, R9S117, R10S118, R10S119, R10S120, R10S121, R10S122, R10S123, R10S124, R10S125, R11S126, R11S127, R11S128, R11S129, R11S130, R11S131, R11S132, R11S133, R11S134, R11S135, R11S136, R11S137, R12S138, R12S139, R12S140, R12S141, R12S142, R12S143, R12S144, R12S145, R12S146

Ji et al. (2012).

Correlation coefficients among the six traits measured in the MGRIL population

Trait I1D I2D I3D I4D CL PL

I1Dz 0.8441** 0.6546** 0.6025** 0.0557 (nsy) 0.5298**
I2D 0.8578** 0.7987** 0.1154 (ns) 0.5264**
I3D 0.9347** 0.3184** 0.4964**
I4D 0.3602** 0.4688**
CL 0.2343**

p < 0.01

I1D=first internode diameter; I2D=second internode diameter; I3D=third internode diameter; I4D=fourth internode diameter; CL=culm length; PL=Panicle length

ns, not significant

Characteristics of QTLs detected for six traits with MGRILs

Traits No. QTL namez Chry Position LODw Additive effectv R2
Interval marker Reference
(cM)x (%)u

First Internode 1 1 27.73 12.56 0.15 21.19 RS014, RM1 sdm1 (Kashiwagi et al. 2008)

2 qI1D1 1 196.62 3.26 -0.07 5.02 RS0124, RS0125

3 qI1D5 5 84.14 6.09 0.10 8.99 RM6841, RS0578

4 6 119.17 3.75 0.08 6.06 RS0693, RM5814 Kashiwagi & Ishimaru (2004)

Second Internode 5 1 27.73 5.86 0.17 12.18 RS014, RM1 sdm1 (Kashiwagi et al. 2008)

6 6 110.87 6.10 0.18 6.31 RS0691, RM1370 Kashiwagi & Ishimaru (2004)

7 7 57.00 3.19 0.13 6.53 STS07015, RS0798 sdm7 (Kashiwagi et al. 2008)

Third Internode 8 qI3D1 1 192.10 3.17 -0.18 6.53 STS01039, RS0124

9 4 74.84 4.12 -0.20 7.76 RS0462, RS0463 qCD-4 (Wang et al. 2011)

10 7 113.39 6.45 0.26 12.94 RM1370, RS0692 Kashiwagi & Ishimaru (2004)

Fourth Internode 11 qI4D1 1 193.10 4.61 0.25 9.97 STS01039, RS0124

12 6 107.87 7.47 0.34 19.25 RS0691, RM1370 Kashiwagi & Ishimaru (2004)

13 12 20.24 3.22 0.19 5.95 RS12140, STS12019 sdm12 (Kashiwagi et al. 2008),

Culm length sd-1 (Ashikari et al. 2002),
Cho et al. 1994,
14 1 171.43 31.14 -9.99 58.23 RS0119, RS0120 Monna et al. 2002,
Sasaki et al. 2002,
(Spielmeyer et al. 2002)

15 qCL5 5 41.08 3.51 2.72 4.24 R5M13, RS0573

16 8 12.16 5.37 3.33 6.44 STS08005, STS08008 pl8 (Xiao et al. 1995),
ph8 (Xiong et al. 1999)

17 qCL12 12 94.2 3.09 2.47 3.57 RS12143, RM1300

Panicle length 18 1 134.68 3.32 0.64 6.59 R1M37, RS0115 pl1 (Xiong et al. 1999),
pl1.1 (Septiningsih et al. 2003)

19 3 41.08 5.44 0.84 11.03 R3M10, RTM5697 Pl3.1 (Septiningsih et al. 2003),
pl3b (Zhuang et al. 1997)

QTLs were newly detected and tentatively named

The number of chromosome

Position of QTL from the top of each chromosome

The logarithm of the ratio of two likelihoods

Positive and negative values indicated additive effect contributed by the alleles of Milyang23 and Gihobyeo, respectively

Percentage of variance explained by each QTL.

Table 1 Result of whole genome re-sequencing of Milyang23 and Gihobyeo

A quality score of 20 represents an error rate of 1 in 100, with a corresponding call accuracy of 99%,

A quality score of 30 represents an error rate of 1 in 1000, with a corresponding call accuracy of 99.9%.

Table 2 Result of read mapping and assembly onto Nipponbare reference genome
Table 3 The list of DNA markers comprising the genetic and physical map constructed in this study

Ji et al. (2012).

Table 4 Correlation coefficients among the six traits measured in the MGRIL population

p < 0.01

I1D=first internode diameter; I2D=second internode diameter; I3D=third internode diameter; I4D=fourth internode diameter; CL=culm length; PL=Panicle length

ns, not significant

Table 5 Characteristics of QTLs detected for six traits with MGRILs

QTLs were newly detected and tentatively named

The number of chromosome

Position of QTL from the top of each chromosome

The logarithm of the ratio of two likelihoods

Positive and negative values indicated additive effect contributed by the alleles of Milyang23 and Gihobyeo, respectively

Percentage of variance explained by each QTL.