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GBS-SNP와 SSR을 이용한 배 ‘만풍배’와 ‘대원홍’ 종간 교배 집단의 고밀도 유전자 연관 지도 작성

김금선1, 김윤경2, 원경호1, 신일섭1, 강삼석1, 김대일3, 김정선4, 정해원1,*

Construction of High-Density Genetic Linkage Map Using GBS-SNPs and SSRs in Interspecific Hybrid Derived from the Cross Between ‘Manpungbae’ and ‘Oharabeni’ Pear (Pyrus spp.)

Korean Journal of Breeding Science 2022;54(4):285-293.
Published online: December 1, 2022

1국립원예특작과학원 배연구소

2국립원예특작과학원 기획조정과

3충북대학교 원예과학과

4국립농업과학원 농업생명자원부 유전체과

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*Corresponding Author: (E-mail: jjhhhwwww@korea.kr, Tel: +82-61-330-1541, Fax: +82-61-330-1503)
• Received: September 2, 2022   • Revised: November 7, 2022   • Accepted: November 7, 2022

Copyright © 2022 by the Korean Society of Breeding Science

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  • Pear (Pyrus spp.) is an economically important fruit tree that grows extensively worldwide. To facilitate the identification of agronomically important traits and provide new information for genetic and genomic research concerning this fruit tree, a high-density genetic linkage map of pear was constructed using 178 F1 populations derived from a cross between ‘Manpungbae’ and ‘Oharabeni’. Single nucleotide polymorphisms (SNPs) detected by genotyping-by-sequencing (GBS) and simple sequence repeats (SSRs) developed from pears were analyzed to construct a genetic linkage map. SSR markers were used to locate the corresponding chromosome number for each linkage group (LG). A total of 1,807 GBS-SNPs and 41 SSRs were anchored to the integrated genetic linkage map. Seventeen LGs were identified, covering a genetic distance of 1,519.4 cM with an average marker density of 0.87 cM. The lengths of the LGs ranged from 70.9 cM (LG 14) to 160.4 cM (LG 15). Each LG had SSR markers from 1 to 5, except for LGs 7, 8, and 9. Our integrated genetic map of pear could be used as a basic frame map for comparative analysis of genomic structure between different pear research groups.
배는 장미과(Rosaceae) 배나무속(Pyrus)에 속하는 온대 과수 작물로 아시아와 유럽 전반에 걸쳐 재배되고 있다. 재배 지역에 따라 서양배와 동양배로 나뉘어지며 동양배는 다시 남방형 동양배와 북방형 동양배로 나뉘어진다(Bell 1990). 배나무속에는 적어도 22개 이상의 기본종들이 포함되어 있다고 보고되어 왔지만 주로 재배되는 종들은 P. pyrifolia, P. ussuriensis, P. bretschneideri, P. communis, P. sinkiangensis로 한정적이며 국내에서는 주로 P. pyrifolia에 속하는 품종들이 재배되거나 품종 육성을 위한 교배친으로 이용되어 왔다(Chung et al. 2020). 최근 소비자들의 소득수준 향상에 따라 과실 품질뿐만 아니라 형태적, 기능적으로 다양한 품종을 요구하고 있어 종간 교배나 특색 있는 유전자원 선발 등을 통한 배 품종들을 육성하고 있다(Peck et al. 2005, Kim et al. 2016). 하지만 배는 영년생 작물로 유년성이 길고, 자가불화합성의 특성을 가지고 있으며, 유전체의 유전적 조성이 복잡하여 육종시 유용 형질과 연관된 유전자를 탐색하는데 긴 시간과 노력이 소요된다. 이는 고밀도 유전자 연관 지도 작성과 양적 형질 유전자좌(Quantitative trait loci; QTL) 분석을 이용한 유용 형질 연관 분자마커 개발로 조기 선발을 통해 육종 연한을 단축시킬 수 있다(Luby & Shaw 2001).
배에서 random amplified polymorphic DNA (RAPD), amplified fragment length polymorphism (AFLP), simple sequence repeats (SSRs) 등 PCR 기반 분자마커를 이용한 유전자 연관 지도 작성에 대한 연구가 보고되어 왔다(Iketani et al. 2001, Dondini et al. 2004, Terakami et al. 2009, Zhang et al. 2013, Yamamoto et al. 2014, Chen et al. 2015). 하지만 이러한 PCR 기반 분자마커는 분석에 많은 시간과 노력이 필요하고 분석 비용이 많이 들기 때문에 고밀도 유전자 연관 지도 구축이나 유용 형질 관련 fine mapping에 적합하지 않다(Wu et al. 2014). 최근에는 PCR 수행 없이도 genotyping을 할 수 있는 single nucleotide polymorphisms (SNPs)를 이용하여 고밀도 유전자 연관 지도를 작성하고 있다.
SNP는 대부분 bi-allele이며 높은 빈도로 변이가 일어나기 때문에 유전체 전반에 고르게 분포하고 있다. 또한 공우성의 특징을 가지고 있으며 재현성이 높아 유전적 구조 분석, 유전적 다양성 확인, QTL 또는 전장유전체연관분석 등 유전 연구에 적합하다(Cho et al. 1999, Gupta et al. 2001, Rafalski 2002, Kujur et al. 2015, Kim et al. 2019, Zhang et al. 2021). Next-generation sequencing (NGS) 기술의 발달은 수십만개의 대용량 SNP를 한번에 탐색 및 genotyping할 수 있게 되었다. 특히 genotyping-by-sequencing (GBS)은 제한효소를 사용하는 sequencing 기술 중 하나로, barcode adapter를 이용해 한번에 많은 샘플들을 sequencing 할 수 있어 다른 NGS 기술들에 비해 비용이 저렴하면서 대용량의 SNP를 얻을 수 있다(Elshire et al. 2011). GBS-SNPs를 이용한 유전 연구는 배뿐만 아니라 사과, 복숭아, 포도 등과 같은 다양한 과수 작물에서 활발히 진행되고 있다(Guo et al. 2019, Kim et al. 2019, Larsen et al. 2019, Thurow et al. 2020).
배의 유전체 해독 연구는 2013년 ‘Dangshansuli’ (P. bretschneideri; Wu et al. 2013)에서 처음 이루어졌으며, 그후 ‘Bartlett’ (P. communis; Chagné et al. 2014)과 ‘Nijisseiki’ (P. pyrifolia; Shirasawa et al. 2021)의 유전체가 해독되었다. 국내에서는 국내 육성 배 품종인 ‘원황’(P. pyrifolia)을 이용하여 유전체 해독이 진행중이다. 배의 유전체 해독이 이루어짐으로써 배 유전 연구에 적합한 많은 수의 분자마커 개발과 더 정밀한 연구가 가능해졌다. 국내 육성 품종이나 육종에 이용되는 교배친은 대부분 P. pyrifolia임에도 불구하고 이전 연구에서는 사과(Malus spp.) 같은 다른 속에서 개발된 SSR 분자마커를 이용하여 유전자 지도가 작성되었으나, ‘원황’ 유전체 기반으로 개발된 SSR 분자마커를 이용한다면 이전보다 더 정확한 유전자 지도 구축이 가능할 것으로 생각된다.
본 연구는 ‘만풍배’(P. pyrifolia)에 ‘대원홍’(P. pyrifolia×P. communis)을 교배한 종간 교배 집단을 이용하여 고밀도 유전자 연관 지도를 구축하기 위해 실시하였다. 유전자 연관 지도 구축에 이용할 SNP 분자마커를 개발하기 위해 GBS를 수행하였다. SSR 분자마커는 ‘원황’(P. pyrifolia) 기반의 SSR 분자마커를 이용하였다.
시험 재료 및 DNA 추출
본 연구의 시험 재료는 Kim et al. (2021)에서 사용된 종간 교배 집단을 이용하여 수행하였다. 교배친은 농촌진흥청 국립원예특작과학원 배연구소에서 보유중인 ‘만풍배’(P. pyrifolia)와 ‘대원홍’(P. pyrifolia×P. communis)을 이용하였다. 교배 모본으로 사용된 ‘만풍배’는 국립원예특작과학원 배연구소에서 1997년에 ‘풍수’(P. pyrifolia)에 ‘만삼길’(P. pyrifolia)을 교배하여 육성된 품종으로 대과종에 속하며 과형은 편원형이고 당도가 높다는 특징이 있다(Cho et al. 2003). 교배 부본인 ‘대원홍’ (P. pyrifolia×P. communis)은 종간 교배 품종으로 ‘만삼길’(P. pyrifolia)에 서양배인 ‘Max Red Bartlett’ (P. communis)을 교배하여 육성되었으며 중소과중의 붉은 과피색이 특징이다(Takahashi et al. 1981). 이 두 품종은 2003년에 국립원예특작과학원 배연구소에서 배 종간 교배 집단의 유전자 연관 지도 작성을 위해 교배되었으며, 이듬해 총 187개의 F1 개체를 획득하였다.
배 종간 교배 집단의 양친인 ‘만풍배’, ‘대원홍’과 F1 187개체중 수세가 안정하고 과실이 착과된 178개체를 선발하여 GBS 분석에 이용하였다. 2017년 5월 양친과 F1 178개체로부터 신초 선단의 어린잎을 채취하여 genomic DNA 추출에 이용하였다. Genomic DNA는 DNeasy Plant Mini Kit (Qiagen, Hilden, Germany)를 이용하여 제조사의 매뉴얼에 따라 추출하였다. 추출된 genomic DNA의 농도와 순도는 각각 DS-11 Spectrophotometer (DeNovix, DE, USA)와 1% agarose gel에 전기영동하여 확인하였다.
GBS 분석 및 SNP 탐색
분리된 ‘만풍배’, ‘대원홍’ 및 F1 178개체의 genomic DNA에 제한효소 ApeKI을 처리한 후 common adapter와 barcode adapter를 ligation하여 GBS 라이브러리를 제작하였다. 제작된 라이브러리는 Illumina Hiseq 2000의 플랫폼을 이용하여 paired-end sequencing을 수행하였으며, 평균 101bp의 reads가 생성되었다. Raw sequencing reads는 barcode sequence에 따라 정렬한 후, SolexaQA v1.13 package를 이용하여 trimming을 수행하였다(Cox et al. 2010). Trimmed reads는 Borrows-Wheeler Aligner (BWA)를 이용하여 배 표준유전체인 P. bretschneideri (‘Dangshansuli’) v1.0 (Wu et al. 2013)의 염기서열에 정렬하고 배 표준유전체에 균일하게 분포하는지 확인하였다(Li & Durbin 2009). Raw SNP 탐색과 consensus sequence 추출은 SAMtools v0.1.16을 이용하여 수행하였다(Li et al. 2009). 그후 ‘만풍배’와 ‘대원홍’의 SNP를 비교 분석하기 위해 SEEDERS in-house script (SEEDERS Inc., Daejeon, Korea)를 이용하여 통합 SNP matrix를 작성하였다. 통합 SNP matrix에서 minor allele frequency (MAF) <5%, missing rate <5%와 같은 필터 기준에 따라 적합하지 않은 SNP를 제외하고 유전자 연관 지도 작성에 이용할 최종 SNP를 선발하였다(Kim et al. 2014).
SSR 분석
교배 모본인 ‘만풍배’와 부본인 ‘대원홍’, F1 178개체에 대해 ‘원황’(P. pyrifolia) 기반으로 개발된 SSR 분자마커 총 320개를 이용하여 유전형 분리비를 확인하였다. 유전형 분리비를 확인하기 위해 양친과 F1 일부를 이용하여 4% 아가로스 겔에서 polymorphic SSR을 1차 선발하였으며, 그 후 양친과 F1 전개체의 유전형 분석을 위해 Shimadzu MCE-202 MultiNA (Shimadzu, Kyoto, Japan)를 이용하여 캐필러리 전기영동을 수행하고 MultiNA Control & Viewer Software (Shimadzu)를 이용하여 유전형 분석을 수행하였다.
유전자 연관 지도 작성
다형성을 나타낸 GBS-SNP와 SSR 분자마커는 JoinMap CP (cross-pollination) code에 따라 교배 모본에서 이형접합을 나타내는 <lm×ll>, 부본에서 이형접합을 나타내는 <nn×np>, 양친에서 모두 이형접합을 나타내는 <hk×hk>, <ef×eg>, <ab× cd>로 전환하였다. 유전자 연관 지도 작성은 JoinMap 4.1 software를 이용하여 수행하였다(Van Ooijen 2006). Linkage group (LG)은 LOD score ≥8을 기준으로 하였고, 분자마커 사이의 연관 거리는 Kosambi's mapping function을 이용하여 계산하였다. 유전자 연관 지도는 MapChart v2.32 프로그램을 이용하여 그림으로 나타내었다(Voorrips 2002).
GBS 및 SNP 탐색
‘만풍배’, ‘대원홍’과 종간 교배 집단 F1 178개체에 대해 GBS를 수행한 결과 총 712.9백만개의 raw sequencing read를 얻었다(Table 1). ‘만풍배’와 ‘대원홍’의 raw sequencing read 수는 각각 4.6백만개, 6.8백만개였고, F1 개체는 평균 4.0백만개의 raw sequencing reads가 생성되었다. Sequencing을 통해 얻은 reads를 trimming한 후, raw sequencing read의 94.5%에 해당하는 총 673.9백만개의 trimmed reads를 얻었다. 개체별 trimmed read수는 845,404개부터 8,096,442개의 범위로 나타났으며, 평균 trimmed read수는 3.8백만개였다. Trimmed reads를 이용하여 배 표준유전체인 P. bretschneideri (‘Dangshansuli’) v1.0의 염기서열에 mapping한 결과, mapping된 reads의 depth는 13.92×로 나타났으며 총 배 표준유전체의 2.76%를 커버하였다. 또한 해독된 염기서열을 바탕으로 ‘만풍배’, ‘대원홍’, F1 178개체의 사이를 비교하여 SNP를 탐색하였다. ‘만풍배’, ‘대원홍’의 종간 교배 집단에서 탐색된 SNP 수는 총 450,350개였으며, 그 중에서 필터링을 통해 최종 29,622개의 SNP를 선발하여 JoinMap 4 수행을 위한 CP code로 전환하였다. CP code에 맞게 genotyping하고 분리비 적합도 결과 유의확률 0.01 미만인 SNP는 제외시켰다. 탐색된 SNP 중 모본에서 이형접합형(<lm×ll>)인 SNP는 2,507개, 부몬에서 이형접합형(<nn×np>)인 SNP는 1,156개, 양친에서 모두 이형접합형(<hk×hk>)인 SNP는 459로 총 4,122개의 SNP가 최종 선발되어 유전자 연관 지도 작성에 이용하였다(Table 2).
SSR 분석
총 320개의 ‘원황’ 기반 SSR중, 122개의 SSR이 ‘만풍배’와 ‘대원홍’의 종간 교배 집단에서 다형성을 나타냈다. 다형성을 보인 SSR 122개 중에 CP code 전환에 적합하지 않은 54개의 SSR을 제거하고 최종적으로 68개를 선발하였다. 최종 선발된 SSR 중 모본에서 이형접합형(<lm×ll>)인 SSR은 10개, 부본에서 이형접합형(<nn×np>)인 SSR는 40개, 양친에서 모두 이형접합형(<hk×hk>, <ef×eg>)인 SSR는 각각 1개와 17개로 전환되었다(Table 2).
유전자 연관 지도 작성
최종 선발된 4,122개의 SNP와 68개의 SSR 분자마커를 이용하여 ‘만풍배’와 ‘대원홍’ 종간 교배 집단 F1 178개체에 분석함으로써 배 유전자 연관 지도를 작성하였다(Fig. 1). ‘원황’ 기반의 SSR 분자마커는 이전 연구에서 염색체 상의 위치가 알려져 있었기 때문에 연관 그룹 번호와 염색체 번호를 일치시킬 수 있었다(Lee et al. 2018). 유전자 연관 지도 작성에 사용된 총 4,190개의 분자마커 중 SNP 1,807개, SSR 41개가 유전자 연관 지도 상에 위치하였다(Table 3). 또한 연관 그룹 7, 8, 9번을 제외한 14개의 연관 그룹에서 적어도 1개 이상의 SSR 분자마커가 위치하였다. SSR 분자마커가 가장 많이 위치한 연관 그룹은 2, 4, 10, 16번으로 각각 5개의 SSR 분자마커가 고정되었다. SSR 분자마커는 시간과 노력, 비용이 많이 들기 때문에 SSR 분자마커만 가지고 고밀도의 유전자 연관 지도를 구축하기엔 어려움이 있다(Piquemal et al. 2005, Wu et al. 2014, Chen et al. 2015). 하지만 SSR 분자마커는 분석이 용이하고 전이성과 재현성이 높을 뿐만 아니라 SNP와 SSR 분자마커를 통합한 유전자 연관 지도는 QTL을 탐색하는데 더 유용하다고 알려져 있기 때문에 본 연구에 이용되었다(Marques et al. 2002, Gasic et al. 2009, Slate et al. 2009). 또한 본 연구에 이용된 ‘원황’ 기반의 SSR 분자마커는 배 유전체 상의 물리적인 위치를 확인할 수 있을 뿐만 아니라 일부 분자마커에서는 유전자의 기능까지 밝혀졌다(Lee et al. 2018).
‘만풍배’와 ‘대원홍’ 종간 교배 집단 유전자 연관 지도의 총 연관 거리는 1,519.4 cM이었으며, 분자마커 간 평균 밀도는 0.87 cM이었다. 길이가 가장 긴 연관 그룹은 169개(SNP 167개, SSR 2개)의 분자마커가 고정된 15번으로 160.4 cM이었다. 배 표준유전체를 구축한 이전의 연구에서도 염색체 15번의 물리적 길이가 가장 길었으며(Xue et al. 2018), 본 연구의 결과와 일치했다. 배에서 다양한 분자마커를 이용하여 유전자 연관 지도를 작성한 연구들이 많이 진행되어왔으며, 본 연구의 유전자 연관 지도의 총 길이는 이전 연구의 유전자 연관 지도의 길이보다 더 길거나 비슷한 수준이었다. Yamamoto et al. (2007, 2014)은 ‘Bartlett’을 이용한 유전자 연관 지도에서는 1,016.1 cM, ‘Taihaku’를 이용한 연관 지도에서는 1,039.1 cM, ‘Akiakari’는 799.1 cM의 연관 지도를 구축하였고, Gabay et al. (2018)은 ‘Spadona’× ‘Harrow Sweet’ 교배 집단에서 1,433.1 cM의 연관 지도를 구축하였다.
최근 차세대 염기서열 분석(next-generation sequencing; NGS)기술이 발달함에 따라 적은 비용에 대용량의 SNP 탐색이 가능해졌다(Zhou et al. 2014, Kujur et al. 2015). SNP 분자마커는 software를 이용하여 SNP 탐색과 동시에 genotyping이 가능하기 때문에 다른 PCR 기반의 분자마커보다 분석이 용이하다(Wang et al. 2017). 따라서 SNP 분자마커를 이용한 유전자 연관 지도는 PCR 기반의 분자마커를 이용한 연관 지도 보다 고밀도의 유전자 연관 지도를 구축할 수 있다. 다양한 작물에서 PCR 기반 분자마커를 이용하여 유전자 연관 지도를 구축해왔으며 평균 밀도는 2.85 cM (Chen et al. 2015), 3 cM (Andargie et al. 2011), 5.1 cM (Wang et al. 2017), 5.6 cM (Tan et al. 2014)으로 낮은 밀도를 보였다. 반면 SNP 분자마커를 이용한 연관 지도의 평균 밀도는 0.56 cM (Zhou et al. 2014), 0.64 cM (Li et al. 2017), 0.98 cM (İpek et al. 2017)으로 PCR 기반 분자마커를 이용한 유전자 연관 지도 보다 고밀도의 유전자 연관 지도가 구축되었다. 따라서 고밀도의 ‘만풍배’와 ‘대원홍’ 종간 교배 집단 유전자 연관 지도를 토대로 QTL 분석을 수행하고 과실 형질 등 유용 형질 연관 유전자를 탐색하여 형질 연관 분자마커를 개발함으로써 배 육종에 도움을 줄 수 있을 것으로 기대된다.
고밀도 유전자 연관 지도 작성을 위하여 종간 교배 집단인 ‘만풍배’(P. pyrifolia), ‘대원홍’(P. pyrifolia×P. communis)과 F1 178개체를 이용하였다. GBS를 통해 개발된 SNP와 ‘원황’ 기반의 SSR을 이용하여 고밀도 유전자 연관 지도를 구축하였다. 총 1,807개의 GBS-SNPs와 41개의 SSR 분자마커가 유전자 연관 지도상에 고정되었으며, 배의 염색체 번호와 일치하는 17개의 연관 그룹이 확인되었다. 유전자 연관 지도의 총 연관 거리는 1,519.4 cM이었으며, 분자마커 간 평균 밀도는 0.87 cM이었다. 가장 작은 연관 그룹의 길이는 연관 그룹 14번의 70.9 cM, 가장 긴 연관 그룹의 길이는 연관 그룹 15번의 160.4 cM이었다. 연관 그룹 7, 8, 9번을 제외한 나머지 연관 그룹에는 최소 1개에서 최대 5개의 SSR 분자마커가 위치하였다. 본 연구에서 구축된 배의 고밀도 유전자 연관 지도를 이용하여 병저항성, 과실 크기, 당 함량 등과 같은 유용 형질 연관 QTL 분석을 수행하고, 이를 통해 관련 유전자를 탐색하거나 분자마커를 개발하여 육종에 이용할 수 있을 것이다. 더 나아가 본 연구에 사용된 전이성 높은 분자마커를 이용하여 장미과내 유전체 비교 분석에 이용할 수 있을 것으로 생각된다.
본 논문은 농촌진흥청 연구사업(세부과제번호 : PJ01358201)의 지원에 의해 이루어진 것임.
Fig. 1
Distribution of GBS-SNP and SSR markers on 17 linkage groups. A black bar indicates a GBS-SNP marker, and a red bar indicates an SSR marker. The left ruler indicates the length in cM.
kjbs-54-4-285-f1a.gifkjbs-54-4-285-f1b.gif
Table 1
Summary statistics of sequencing reads generated by genotyping-by-sequencing.
Table 1
Parameter Data
Total number of raw sequencing reads 712,909,282
Total number of trimmed reads 673,909,914 (94.53%)
Total length of trimmed reads 56,576,458,887 bp
Total number of mapped reads 513,019,876 (76.13%)
Total of number of mapped regions 15,176,795
Average depth of mapped region 13.92 ×
Total length of mapped regions 2,518,884,203 bp
Reference genome coverage 2.76%
Table 2
A total number of SNP and SSR markers according to segregation types used for construction of genetic linkage map.
Table 2
Segregation typez No. of SNPs No. of SSRs
hk×hk 459 1
ef×eg - 17
lm×ll 2,507 10
nn×np 1,156 40
Total 4,122 68

zhk×hk and ef×eg, heterozygous in both maternal parent and paternal parent; nn×np, homozygous in maternal parent and heterozygous in paternal parent; lm×ll, heterozygous in maternal parent and homozygous in paternal parent

Table 3
Summary of the high-density genetic map constructed using an F1 individuals derived from the cross of ‘Manpungbae’ and ‘Oharabeni’.
Table 3
Linkage group No. of SNPs No. of SSRs Total genetic distance (cM) Average genetic distance (cM) Max. gap (cM) <5 cM gap
1 69 3 71.3 0.99 8.17 69
2 55 5 87.1 1.45 12.59 57
3 106 2 83.3 0.77 6.74 104
4 113 5 77.7 0.66 6.24 116
5 138 1 99.2 0.71 4.42 139
6 95 1 74.8 0.78 3.63 96
7 144 - 85.0 0.59 4.36 144
8 60 - 79.4 1.32 13.17 58
9 123 - 78.7 0.64 3.69 123
10 125 5 104.5 0.80 11.52 127
11 113 4 88.4 0.76 12.79 114
12 115 2 77.8 0.66 4.56 117
13 74 2 85.6 1.13 6.57 75
14 128 2 70.9 0.55 4.07 130
15 167 2 160.4 0.95 9.39 164
16 98 5 91.6 0.89 13.55 100
17 84 2 103.7 1.21 11.46 82
Total 1,807 41 1,519.4 13.55 1,815
Average 0.87
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Construction of High-Density Genetic Linkage Map Using GBS-SNPs and SSRs in Interspecific Hybrid Derived from the Cross Between ‘Manpungbae’ and ‘Oharabeni’ Pear (Pyrus spp.)
Korean. J. Breed. Sci.. 2022;54(4):285-293.   Published online December 1, 2022
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Construction of High-Density Genetic Linkage Map Using GBS-SNPs and SSRs in Interspecific Hybrid Derived from the Cross Between ‘Manpungbae’ and ‘Oharabeni’ Pear (Pyrus spp.)
Korean. J. Breed. Sci.. 2022;54(4):285-293.   Published online December 1, 2022
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Construction of High-Density Genetic Linkage Map Using GBS-SNPs and SSRs in Interspecific Hybrid Derived from the Cross Between ‘Manpungbae’ and ‘Oharabeni’ Pear (Pyrus spp.)
Image
Fig. 1 Distribution of GBS-SNP and SSR markers on 17 linkage groups. A black bar indicates a GBS-SNP marker, and a red bar indicates an SSR marker. The left ruler indicates the length in cM.
Construction of High-Density Genetic Linkage Map Using GBS-SNPs and SSRs in Interspecific Hybrid Derived from the Cross Between ‘Manpungbae’ and ‘Oharabeni’ Pear (Pyrus spp.)

Summary statistics of sequencing reads generated by genotyping-by-sequencing.

Parameter Data
Total number of raw sequencing reads 712,909,282
Total number of trimmed reads 673,909,914 (94.53%)
Total length of trimmed reads 56,576,458,887 bp
Total number of mapped reads 513,019,876 (76.13%)
Total of number of mapped regions 15,176,795
Average depth of mapped region 13.92 ×
Total length of mapped regions 2,518,884,203 bp
Reference genome coverage 2.76%

A total number of SNP and SSR markers according to segregation types used for construction of genetic linkage map.

Segregation typez No. of SNPs No. of SSRs
hk×hk 459 1
ef×eg - 17
lm×ll 2,507 10
nn×np 1,156 40
Total 4,122 68

Summary of the high-density genetic map constructed using an F1 individuals derived from the cross of ‘Manpungbae’ and ‘Oharabeni’.

Linkage group No. of SNPs No. of SSRs Total genetic distance (cM) Average genetic distance (cM) Max. gap (cM) <5 cM gap
1 69 3 71.3 0.99 8.17 69
2 55 5 87.1 1.45 12.59 57
3 106 2 83.3 0.77 6.74 104
4 113 5 77.7 0.66 6.24 116
5 138 1 99.2 0.71 4.42 139
6 95 1 74.8 0.78 3.63 96
7 144 - 85.0 0.59 4.36 144
8 60 - 79.4 1.32 13.17 58
9 123 - 78.7 0.64 3.69 123
10 125 5 104.5 0.80 11.52 127
11 113 4 88.4 0.76 12.79 114
12 115 2 77.8 0.66 4.56 117
13 74 2 85.6 1.13 6.57 75
14 128 2 70.9 0.55 4.07 130
15 167 2 160.4 0.95 9.39 164
16 98 5 91.6 0.89 13.55 100
17 84 2 103.7 1.21 11.46 82
Total 1,807 41 1,519.4 13.55 1,815
Average 0.87
Table 1 Summary statistics of sequencing reads generated by genotyping-by-sequencing.
Table 2 A total number of SNP and SSR markers according to segregation types used for construction of genetic linkage map.

zhk×hk and ef×eg, heterozygous in both maternal parent and paternal parent; nn×np, homozygous in maternal parent and heterozygous in paternal parent; lm×ll, heterozygous in maternal parent and homozygous in paternal parent

Table 3 Summary of the high-density genetic map constructed using an F1 individuals derived from the cross of ‘Manpungbae’ and ‘Oharabeni’.