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배 ‘원황’() 유전체 해독에 기반한 SSR 마커 개발 및 유전자 지도 작성

이지윤1,2, 서미숙1, 원소윤1, 임경아1, 신일섭3, 최동수2,*, 김정선1,*

Construction of a Genetic Map using the SSR Markers Derived from “Wonwhang” of Pyrus pyrifolia

Ji Yun Lee1,2, Mi-Suk Seo1, So Youn Won1, Kyoung Ah Lim1, Il Sheob Shin3, Dongsu Choi2,*, Jung Sun Kim1,*
Korean Journal of Breeding Science 2018;50(4):434-441.
Published online: December 1, 2018

농촌진흥청 국립농업과학원 농업생명자원부 유전체과

군산대학교 생물학과

농촌진흥청 국립원예특작과학원

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* (Jung Sun Kim, E-mail: jsnkim@korea.kr, Tel: +82-63-238-4559, Fax: +82-63-238-4552)
* (Dongsu Choi, E-mail: choid@kunsan.ac.kr, Tel: +82-63-469-4583, Fax: +82-63-469-7421)
• Received: September 12, 2018   • Revised: October 30, 2018   • Accepted: November 9, 2018

Copyright: © 2018 by the Korean Society of Breeding Science

This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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  • High-density genetic linkage mapping is critical for undertaking marker-assisted selection and confirming quantitative trait loci, as well as helping to build pseudomolecules of genomes. We constructed a genetic map using 94 F1 populations generated from the interspecific cross between Korean cultivar “Wonwhang” (Pyrus pyrifolia, NCBI BioSample SAMN05196235) and European cultivar “Bartlett” (Pyrus communis). We designed a total of 24,267 SSR markers based on the genome sequences of “Wonwhang” for this. To select the markers that are linked to the traits important in pear breeding programs, SSR-containing genomic sequences were subjected to nucleotide sequence homology searches, which resulted in 510 SSR markers with high similarity to genes encoding proteins with putative functions such as transcription factors, resistance proteins, flowering time, and regulatory genes. Of these, 70 markers showed polymorphisms in parents and segregating populations and were used to construct a genetic linkage map, together with the unpublished 579 SNPs obtained from genotyping by sequencing analysis. The genetic linkage map covered 3,784.2 cM and the average distance between adjacent markers was 5.8 cM. Seventy SSR markers were distributed across 17 chromosomes with more than one locus.
  • Keywords: Pear; Pyrus pyrifolia; SSR marker; Linkage map
배(Pyrus spp.)는 포도(Vitis), 사과(Malus), 핵과류(Prunus)와 함께 중요한 온대 과수로, 장미과(Rosaceae) 배나무아과(Maloideae) 배나무속(Pyrus)에 속하는 다년생 식물이다(Bell 1991; Wu et al. 2013). 사과, 배와 같이 배나무아과에 속하는 식물들은 장미과 원시조상(n=9)으로부터 Whole genome duplication (WGD)이 일어난 후, 염색체의 융합 혹은 결실되는 과정을 통해 17개 염색체(2n=34)를 가지고 있다. 반면 복숭아, 딸기 같은 다른 장미과 식물들은 WGD이 일어나지 않았다(Ro owley 1967; Wu et al. 2013). 다년생 식물인 배는 긴 유년기를 가지고 있고, S 유전자좌에 의해 조절되는 자가불화합성(self-incompatibility)으로 인하여 높은 수준의 이형 접합을 보임으로써, 특정 형질을 조절하는 유전자를 발굴하거나, 새로운 품종 개발을 위한 육종연구에 많은 시간과 노력이 필요하다(Chen et al. 2015; Wu et al. 2014; Yamamoto & Chevreau 2009). 따라서 초기 세대의 어린 단계에서 작물을 선발하는 분자 마커 이용 선발법(maker- Assisted Selection, MAS)을 위한 분자 마커 개발과, 유전자 연관 지도를 작성하여 양적 형질 유전자좌(quantitative trait loci, QTLs)를 확인하는 연구는 배의 육종연구를 위해 매우 중요하다고 할 수 있다(Tanksley et al. 1989). 분자 마커는 표현형의 특징을 비교하는 형태 마커(morphological marker)와는 다르게 DNA 염기서열의 차이를 기반으로 하기 때문에 많은 양의 시료 분석이 가능할 뿐만 아니라, 작물의 생육 변화 및 환경적 요인에 크게 영향을 받지 않는다(Collard et al. 2005). 이러한 이유로 식물 육종 분야에서는 유전적 특성을 탐구하기 위한 분자기술의 개발과 함께 다양한 분자 마커가 개발되고 있다. 배나무속에서는 RFLP (Restriction Fragment Length Polymorphism) (Monte-Corvo et al. 2000; Yamamoto et al. 2007), RAPD (Random Amplified Polymorphic DNA) (Yuanwen Teng et al. 2001; Yuying et al. 2006), AFLP (Amplified Fragments Length Polymorphism) (Dolatowski et al. 2004; Zhang et al. 2013), SRAP (Sequence-related amplified polymorphism) (Zhao et al. 2013), SSR (Simple Sequence Repeat) (Chen et al. 2015; Fan et al. 2013)과 같은 다양한 분자 마커의 개발을 통해 유전자 지도 작성, 계통 유연 관계(phylogenetic relationship) 분석, 유전적 다양성 분석, 그리고 특정 형질과 관련된 QTL 연구가 진행 중에 있다. 최근 SNPs 활용 유전지도 작성이 이루어졌는데, Saeed et al. (2013)은 유럽 배(P. communis)의 857개 SNPs 활용 유전지도를, Terakami et al. (2014)은 일본 배 ‘Hosui’에 대한 609개 SNPs가 위치한 유전지도를 보고하였다.
SSR 마커의 경우, 진핵 생물의 유전체에 다수 존재하는 2~10 bp의 짧은 염기서열의 반복에 기반을 두어 개발된 마커로, PCR 분석과 같은 손쉬운 방법으로 높은 다형성(polymorphism)과 재현성(reproducibility)을 가지는 공우성(co-dominant) 마커를 탐색할 수 있기 때문에 많은 식물에서 분자 마커로 활용되고 있다(Kimura et al. 2002, Yamamoto et al. 2002a). 반면에, SSR 마커의 개발을 위해서는 분석하고자 하는 작물의 염기서열 정보와 함께, 대립 유전자들의 증폭을 위한 특정 primer 제작에 많은 초기 비용이 소모되는 단점을 가지고 있다(Pierantoni et al. 2004). 최근, 차세대 염기서열 분석(Next Generation Sequencing, NGS) 기술의 발전으로 다양한 식물체에서 표준 유전체 정보가 해독되고 있으며, Wu et al. (2013)에 의해 북방형 동양 배(중국 배) 유전체 정보가 밝혀짐으로써, SSR 마커를 포함한 다양한 분자 마커들의 개발이 활발하게 진행 중에 있다(Chen et al. 2015; Yamamoto et al. 2002a; Yamamoto et al. 2002b).
배의 유전체 해독연구는 중국배(P. bretschneideri Rehd.) ‘Danshansuli’와, 서양배(P. communis L.) ‘Bartlett’에서 이루어졌으나(Chagné et al. 2014; Wu et al. 2013), 최근 대만 배, 일본 배 등 남방형 동양 배에 대하여 Illumina sequencing에 의한 draft genome이 보고되었다(Takameura et al. 2018a, b). 유전자 영역에 대한 예측 및 해석이 없는 조립한 게놈 시퀀스 보고로서 유전체 해독연구는 아니다. 한국 배에 대한 보고로서, Oh et al.(2015)은 황금배와 미내배에 대한 36.64X, 31.04X 각 각 NGS 분석하고, 중국배 ‘Danshansuli’ 유전체에 비교 맵핑하여 다양한 다형성 마커를 보고하였다. 농촌진흥청에서는 남방형 동양 배에 속하는 ‘원황’의 유전체 해독 연구를 진행중이다. 배 ‘원황’ 게놈 해독은 장거리 염기서열분석을 수행하고, 이형접합인(hetrosequences) 염기서열을 보정한 후, 3종의 mate-paired 라이브러리 활용 조립하여 2,221개 scaffold를 조립하였다(미보고). 조립된 scaffold의 염색체 위치를 알고자 Genotyping-by- sequencing (GBS) 분석을 수행하여 유전자지도를 작성하였다. 본 유전지도에 SSR 마커들을 위치시켜, 과실의 주요 형질(단맛, 신맛), 개화기, 과실 특성에 대한 QTL을 분석하고, QTL이 위치한 영역의 SSR을 선발하여 육종프로그램을 지원하고자 수행하였다. 반복서열 데이터베이스에서, SSR motif를 분석하여 24,267개의 SSR 프라이머 조합을 제작하였다. 탐색된 SSR motif가 포함되어 있는 시퀀스 정보의 유전자 상동성 분석(BLASTn)을 수행하고, 전사인자나 병 저항성 유전자, 개화 관련 유전자 및 대사조절유전자 등과 상동성이 높은 서열을 가지고 있는 잠재적 유용유전자를 탐침하는 SSR 마커 510개를 선발하였다. 이들 SSR 마커들을 활용하여 양친과 분리집단에서 다양성을 보이는 88개 마커들의 분리비를 활용 유전자지도를 작성하였다.
식물 재료
본 실험에서는 농촌진흥청 국립원예특작과학원에서 제공받은 F1 분리집단을 대상으로 분석을 수행하였다. 남방형 동양배 ‘원황’(Pyrus pyrifolia)을 모본, 서양배 ‘Bartlett’ (Pyrus communis)을 부본으로 사용하여 종간 교잡한 F1 분리 집단 191개체에서 생육이 양호한 94개체를 선발하여 실험에 사용하였다.
Genomic DNA 추출
양친과 F1 교배집단 94 개체에서 만개 20일 후의 어린잎을 채취하여 액체질소를 이용하여 조직을 마쇄한 후, DNeasy Plant Mini Kit (Qiagen, Germany)의 매뉴얼에 따라 genomic DNA를 추출하였다. 추출된 DNA는 1% agarose gel에서 전기영동 하여 DNA를 확인하였고, nanodrop 1000 spectrophotometer (Thermo Scientific, USA)를 이용하여 농도 및 순도를 확인한 후, 1 µl당 5 ng의 농도로 희석하여 PCR 분석에 이용하였다.
SSR 마커를 이용한 PCR 분석
배의 유전자 지도 작성에 사용할 SSR primer는 ‘원황’ scaffold의 반복서열에서 제작된 24,267개의 SSR primer 중에서 di-nucleotide motif를 가진 3,200개를 선발하였다. SSR 마커가 포함되어 있는 게놈 시퀀스들을 선별하고자, NCBI 뉴클레오타이드 데이터베이스와 상동성 분석을 E-value≤1e-30 조건에서 수행하였다. 추정되는 기능이 전사인자(transcription factor), 병 저항성 유전자, 개화기 등 조절 유전자 등 주요한 기능이 있는 유전자를 증폭하기 위한 PCR 프라이머 조합을 선별하였다. 선발된 primer로 양친과 ‘원황’ × ‘Bartlett’ 분리집단 94개체를 대상으로 PCR 분석을 수행하였다. PCR 분석은 30 ng genomic DNA와 10 pmol/µl의 양방향 프라이머를 넣어 12 µl를 맞추고, 2배 농도의 TOPsimple™DyeMIX-nTaq (Taq DNA polymerase 0.2 U, dNTPs 0.4 mM, MgCl2 3 mM; Enzynomics, Daejeon, Korea)을 12 µl 넣은 조성액을, 반복 없이, 94°C에서 5분 동안 pre-denaturation을 수행한 뒤, 94°C에서 30초, 60°C에서 30초, 72°C에서 1분의 조건을 30회 반복하는 조건으로 수행하였다. 일부 primer의 경우 Tm 값에 따라 온도를 조정하였다. PCR 산물은 2.5~3% agarose gel을 만든 후, 1× TAE (Tris–Acetate EDTA) buffer내에서 80 V에 3시간 30분 동안 전기영동 하여 양친과 분리집단 다양성을 탐색하였다.
분리비 판독 및 유전자 지도 작성
SSR primer는 모본(원황)과 부본(Bartlett)의 대립 단편(allele) 증폭 양상에 따라 모본이 헤테로인 경우는 모본계 마커(WH), 부본이 헤테로인 경우에는 부본계 마커(BT), 모본과 부본이 모두 헤테로인 경우에는 양친 헤테로 마커(WB)로 구분하였다. 본 연구에서 사용한 모든 마커들은 각 마커들이 만들어진 scaffold와 위치를 나타낼 수 있도록 명명하였다. 예를 들어 WH_W1325_101 마커는 원황 1325번 스캐폴드 시퀀스의 101Kb에서 만들어진 마커를 말한다. SSR 마커들은 헤테로마커 구분 표시 뒤에(WH, BT, WB) ‘SSR’ 표기하여 구분되도록 하였다. 양친의 분리비와 분리집단 분리비를 JoinMap 4.1에서 기술한 바 대로 판독하였다. SSR 마커의 다형성을 조사하기 위하여 마커별 다형성 정도를 나타내는 지수인 polymorphic information content (PIC) 값을 Anderson et al. (1993)의 공식을 이용하여 산출하였다. 유전자지도 작성은 JoinMap4.1 (Kyazma B.V., Wageningen, Netherlands)을 이용하였고, independence LOD값 2~10 조건과, 유사도 threshold는 0.95 이상으로 분석하였다.
양친 및 분리집단 SSR 분석
‘원황’의 2,221개 scaffold에서 반복서열을 추출하고, SSR을 제놈와이드 탐색하였다. SSR이 포함된 영역을 추출하고, Primer3 프로그램 활용 프라이머를 디자인 하였다(Koressar & Remm 2007). SSR 반복서열이 AC/AG/AT/CG와 같이 di-nucleotide가 87.11%, AAT/AAG/AAC/ATG/AGT/AGG/AGC 등 tri-nucleotide가 9.66%로서, di-와 tri-nucleotide가 전체의 96.77%가 분포하였다. 탐색한 SSRs을 포함하는 유전체 서열을 추출하고 이를 PCR 검출할 수 있는 24,267개 쌍의 SSR 프라이머를 디자인하였다. SSR 모티브 유형에 따라, di-nucleotide에서 deca-nucleotide까지 아홉 개 종류의 반복서열 타입에 따라 분류한 결과는 Table 1과 같다. ‘원황’ scaffold 2,221개 중 우선적으로 1번부터 842번까지 scaffold 반복시퀀스에서 3,200개 SSR 마커가 디자인되었다. 이들 마커가 만들어진 시퀀스들에 대하여 NCBI DB와 염기서열 상동성 분석(BLASTn)을 하여, SSR 마커들이 유래된 서열정보가 전사인자나 병 저항성 유전자, 개화 관련 유전자 및 대사조절유전자 등과 상동성이 있는 마커들을 선발하였다. 총 510개 SSR 마커쌍을 선별하고 프라이머를 합성하였다. 그 예를 보면, auxin-induced protein 15A (LOC103454085), tobamovirus multiplication protein 1 (LOC103948155), TMV resistance protein N (LOC103935600), serine carboxypeptidase II-3 (LOC103957119), heat stress transcription factor B-3 (LOC103931654), cytochrome P450 86B1 (LOC103403397) 등이다. 양친과 그 F1 분리집단 94개에 대하여, 선발한 510개 프라이머쌍으로 PCR 분석하여 2.5~3.0% agarose gel상에서 전기영동하였다, 양친 및 F1 분리집단에서 다형성을 보인, 88개의 마커(Supplementary Table 1)에 대한 분리비를 판독하였는데, 이들 마커들은 25개의 모본계 마커(WH), 53개의 부본계 마커(BT), 10개의 양친 헤테로 마커(WB)로 분류되었다.
Table 1
Classification to the repeat types, frequency and number of primers derived from Pyrus pyrifolia cv. Wonwhang genome sequence.
Table 1
Repeat type Frequency Primer pair (%)
Di-nucleotide 102,797 21,200 20.62
Tri-nucleotide 11,403 2,695 23.63
Tetra-nucleotide 2,991 458 15.31
Penta-nucleotide 454 71 15.64
Hexa-nucleotide 203 41 20.2
Hepta-nucleotide 137 8 5.84
Octa-nucleotide 16 3 18.75
Ennea-nucleotide 0 0 0
Deca-nucleotide 2 0 0
선발된 SSR 마커를 기반으로 한 유전자 지도 작성
GBS 분석을 통해 SNP 마커가 포함된 유전자지도에 88개 SSR마커를 연계하여 JoinMap4.1 프로그램으로 분석하였다. 분석한 결과, 배 염색체 17의 염색체 상에 70개의 SSR 마커가 위치하였다. 염색체 당 최소한 한 개 이상의 SSR 마커가 위치하였다(Fig. 1). 총 유전거리는 3784.17cM이고, 마커들 사이의 평균거리는 5.8cM 이었다(Table 2). 유전자지도에 위치하는 SSR 마커에 의해 분석된 대립 단편(allele)의 수는 2~5개였고, 총 205개가 나타났으며, 마커 당 평균 2.93개의 대립 단편을 확인하였다(Supplementary Table 2). 또한 F1 집단의 다형성 정도를 조사한 결과, 각 마커 별 다형성 정도를 나타내는 지수인 PIC값은 0.346에서 0.769의 범위로 나타났으며, 평균값은 0.54이었다. SNP 마커들과 SSR 마커들의 구분을 위하여 SSR 마커들은 파란색으로 표기하였다(Fig. 1). 모든 마커들은 마커들이 유래한 scaffold를 표시하였는데, W0085는 원황 scaffold 85번이고, 그 길이는 1.78 M이다. 이 scaffold의 위치는 Chr1 65.5cM에, WH_W0085_301 SNP 마커로 4.3cM 인접한 BT_SSR_W0085_0135, SSR 마커로 맵핑되었다. SSR마커들은 같은 스캐폴드에 SNP 마커들과 잘 인접되어 맵핑되었다. 이들 scaffold 맵핑 정보들은 ‘원황’의 Pseudomolecule 작성에 활용할 예정이다.
Table 2
Integrated genetic map of SSRs in Pyrus pyrifolia.
Table 2
Molecular Marker Chromosome number Total
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Loci
SSR 3 9 3 4 7 3 3 1 7 8 4 1 3 1 5 6 2 70
SNP 18 38 40 27 43 36 22 27 22 31 36 43 45 30 57 30 34 579
Total Marker 21 47 43 31 50 39 25 28 29 39 40 44 48 31 62 36 36 649
Length (cM) 189.6 361.1 218.5 213.2 169.8 279.9 140.7 132.8 171.3 257.3 235.9 277.5 341.5 218.3 320.1 96.8 159.7 3784.2
Average Interval 9.0 7.7 5.1 6.9 3.4 7.2 5.6 4.7 5.9 6.6 5.9 6.3 7.1 7.0 5.2 2.7 4.4 5.8
Fig. 1
Integrated genetic map constructed with 70 SSR unpublished 579 SNP markers in Pyrus pyrifolia. This genetic map covers 3,784.2cM and the average distance between makers is 5.8cM
KJBS_50_434_fig_1.jpg
유전자 지도에 맵핑된 SSR 마커가 위치한 시퀀스의 추정되는 기능들은 phloem protein 2 (PP2)로 알려진 PHLOEM PROTEIN 2-LIKE A10 (WH_SSR_W0144_0368, Chr02, 36.8cM) (Gómez & Pallás 2001), 식물 호르몬, 리그닌, 플라보노이드등의 생합성에 관여하는 효소인 cytochrome P450 86B1 (WH_SSR_W0103_1927, Chr02, 361cM) (Watson et al. 2001), 그리고 식물 병저항성 관련 유전자, TMV resistance protein N (BT_SSR_W1584_1671, Chr11, 82.5cM) (Dinesh-Kumar et al. 2000) 이었다(Supplementary Table S2). 본 연구에서는 유용한 70개 SSR 마커들의 유전위치를 확인할 수 있었고, 이들 SSR 마커들은 향후 배 연구에서 양적 형질 유전자좌(QTLs)의 위치를 예측하거나 배 형질 개선을 위한 유전 육종에 마커로 활용될 수 있으리라 기대한다.
본 연구에서는 배 ‘원황’(Pyrus pyrifolia)의 유전체 정보를 바탕으로, 유용 유전자 관련 SSR 마커를 선발하였고, 선발된 SSR과 SNP 마커를 이용하여 ‘원황’ × ‘Bartlett’ F1 교배집단에 대한 유전자 지도를 작성하였다. ’원황’의 scaffold에서 제작된 SSR 마커 유래 염기서열들과 NCBI nucleotide DB와 BLASTn 분석하여, 유용한 유전자들과 높은 상동성을 보이는 510개 SSR 마커를 선발하였다. 이들 마커를 사용하여 양친과 F1 집단 94개체의 대립 단편의 증폭 양상을 확인한 결과, 88개 마커들이 헤테로 집단에 맞는 분리비를 보였다. 선발된 88개의 SSR 마커는 GBS 분석을 통해 획득한 579개 SNP 마커와 함께 ‘원황’의 유전자지도를 작성하였다. 70개의SSR 마커들은 배 염색체 수와 같은 17개의 염색체에 잘 위치하였고, 모든 염색체에 한 개 이상의 마커로 위치하였다. 유전자지도의 총 유전거리는 3784.2cM이고 마커간 평균거리는 5.8cM이었다. 본 연구에서 개발된 SSR 분자 마커 및 이를 기반으로 만들어진 유전자지도는 배의 육종 및 유전 연구에 유용한 정보를 제공할 것으로 기대한다.
본문의 Supplementary Table 1과 Supplementary Table 2는 한국육종학회지 홈페이지에서 확인할 수 있습니다.
본 연구는 농촌진흥청 국립농업과학원 어젠다 사업(과제번호: PJ013381022018, PJ010450012017)의 지원에 의하여 수행되었습니다.
KJBS-50-434_suppl.pdf
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Construction of a Genetic Map using the SSR Markers Derived from “Wonwhang” of Pyrus pyrifolia
Korean. J. Breed. Sci.. 2018;50(4):434-441.   Published online December 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.4.434
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Construction of a Genetic Map using the SSR Markers Derived from “Wonwhang” of Pyrus pyrifolia
Korean. J. Breed. Sci.. 2018;50(4):434-441.   Published online December 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.4.434
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Construction of a Genetic Map using the SSR Markers Derived from “Wonwhang” of Pyrus pyrifolia
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Fig. 1 Integrated genetic map constructed with 70 SSR unpublished 579 SNP markers in Pyrus pyrifolia. This genetic map covers 3,784.2cM and the average distance between makers is 5.8cM

Fig. 1

Construction of a Genetic Map using the SSR Markers Derived from “Wonwhang” of Pyrus pyrifolia

Classification to the repeat types, frequency and number of primers derived from Pyrus pyrifolia cv. Wonwhang genome sequence.

Repeat type Frequency Primer pair (%)
Di-nucleotide 102,797 21,200 20.62
Tri-nucleotide 11,403 2,695 23.63
Tetra-nucleotide 2,991 458 15.31
Penta-nucleotide 454 71 15.64
Hexa-nucleotide 203 41 20.2
Hepta-nucleotide 137 8 5.84
Octa-nucleotide 16 3 18.75
Ennea-nucleotide 0 0 0
Deca-nucleotide 2 0 0

Integrated genetic map of SSRs in Pyrus pyrifolia.

Molecular Marker Chromosome number Total
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Loci
SSR 3 9 3 4 7 3 3 1 7 8 4 1 3 1 5 6 2 70
SNP 18 38 40 27 43 36 22 27 22 31 36 43 45 30 57 30 34 579
Total Marker 21 47 43 31 50 39 25 28 29 39 40 44 48 31 62 36 36 649
Length (cM) 189.6 361.1 218.5 213.2 169.8 279.9 140.7 132.8 171.3 257.3 235.9 277.5 341.5 218.3 320.1 96.8 159.7 3784.2
Average Interval 9.0 7.7 5.1 6.9 3.4 7.2 5.6 4.7 5.9 6.6 5.9 6.3 7.1 7.0 5.2 2.7 4.4 5.8
Table 1 Classification to the repeat types, frequency and number of primers derived from Pyrus pyrifolia cv. Wonwhang genome sequence.
Table 2 Integrated genetic map of SSRs in Pyrus pyrifolia.

Table 1

Table 2

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