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Korean. J. Breed. Sci. : Korean Journal of Breeding Science

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ARTICLE

Genetic Diversity of Wild Pear Accessions Collected in Korea


Published online: February 28, 2015

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*Corresponding author (E-mail: yskwon3@dau.ac.kr, Tel: +82-51-200-7521, Fax: +82-51-200-7505)
• Received: February 12, 2015   • Revised: February 18, 2015   • Accepted: March 2, 2015

© The Korean Society of Breeding Science

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  • The objective of this study was carried out to assess genetic diversity of Korean local accessions of wild pear using morphological traits and microsatellite markers. Assessment of 14 phenotypic traits showed high variation among wild pear accessions. These parameters were not applicable genetic diversity analysis of wild pear collection due to quantitatively inherit and their expression is affected by environmental factors. Microsatellite markers were used to evaluate the genetic similarity of 62 accessions. Among 50 tested SSR markers, 16 primer pairs were selected to profile genetic diversity on the basis of high level polymorphism. These microsatellite markers showed 1 or 2 discrete amplified fragment for all of the accessions. The sixteen microsatellite loci amplified 278 alleles, with 10 to 27 alleles (average 17.385) per locus. The mean of observed heterozygosity and polymorphism information content were 0.776 and 0.836, respectively. The Jaccard’s similarity coefficient ranged from 0.09 to 1.00. Two major groups were produced from all the accessions by UPGMA cluster analysis. Fifty five accessions could be discriminated except for 2 pairs. These results would be provided useful information about valuable genetic resources though assessment of genetic diversity and relationship in local pear accessions.
돌배나무(Pyrus pyrifolia)는 장미과에 속하는 목본성 다년 생 식물로 우리나라, 일본, 중국 등 동북아시아 지역에 주로 자생하고 있다. 우리나라의 경우 전국 산간지역에 주로 분포 하며 내한성이 강하고 순백색의 꽃이 화려하기 때문에 도시 공원의 경관수로 활용 가치가 크고, 열매는 한약재료로 활용 도가 증가되고 있는 수종의 하나이다. 배나무는 유럽, 미국, 아 프리카에서 재배되는 서양배나무(P. communis L.), 중국에서 유래된 북방형 돌배나무(P. bretschneideri Rehd., P. ussuriensis Maxim.), 우리나라와 일본에서 자생하고 재배되고 있는 남방 형 돌배나무(P. pyrifolia Nakai) 로 크게 나눌 수 있다(Bell, 1990). 우리나라에는 배나무 속 10종, 17변종, 10품종의 야생 또는 재배종이 분포하고 있으며, 일본의 남방형 돌배나무의 경 우 중국과 한반도가 기원인 것으로 알려지고 있다(Shimamura 1998).
유전자원의 특성 평가는 주로 형태적 형질 조사를 활용하 고 있는데 돌배나무와 같은 목본류 식물은 수령에 따른 잎, 꽃, 과실 등과 같은 형질에서 오차가 발생할 가능성이 있기 때문에 수집 유전자원의 정확한 특성 평가는 단순하지 않다. 또한, 다수의 유전자가 관여하는 양적 형질은 환경에 의해 영 향을 받기 때문에 조사 형질의 연차간 변이 등과 같은 문제점 을 내포하고 있으며, 형태적으로 유사한 형질을 가진 유전자 원 및 품종을 구별하기가 쉽지 않다. 이러한 문제점을 해결할 수 있는 방법이 분자표지 인자에 의한 유전자원의 특성 평가 방법이다. 분자표지에 의한 배 유전자원에 대한 특성 방법은 Inter-simple sequence repeat (ISSR) (Zhu et al. 2009), Amplified fragment length polymorphism (AFLP) (Cho et al. 2009), microsatellite (Miranda et al. 2010, Rugienius et al. 2013, Sehic et al. 2012) 마커가 활용되고 있다. 최근에는 반복 재현성이 높고 유전자원내에 다형성 정도가 높은 microsatellite 방법을 주로 활용하고 있다.
Microsatellite 마커를 이용한 배나무에 대한 연구는 일본 에서 가장 활발하게 이루어져 있는데 일본배나무와 서양배나 무의 종간 교잡을 통해 양성된 집단을 활용한 유전자 지도 작 성(Yamamoto et al. 2002), Simple sequence repeat (SSR) 마커를 이용한 품종 식별(Kimura et al. 2002), 배나무 과일 과 가공품(통조림, 주스, 건조된 과일)에 대한 DNA 분석 방 법 개발(Yamamoto et al. 2006), 동양배나무에서 유래된 Expressed sequence tag (EST)로 부터 새로운 SSR 마커의 개발(Nishitani et al. 2009) 등과 같은 연구 결과가 보고되고 있다. 배나무 유전자원 특성 평가에 microsatellite 마커를 사 용한 예는 중국(Bao et al. 2007), 스페인(Miranda et al. 2010), 폴란드(Wolko et al. 2010), 스웨덴(Sehic et al. 2012) 등과 같은 국가에서 자국에서 수집된 유전자원 및 품종을 대상으 로 8~17개의 마커를 사용하였다는 점이다. 우리나라에서는 농촌진흥청 국립원예특작과학원에서 AFLP 마커를 활용하여 일본, 중국, 미국 등에서 수집된 배나무 유전자원의 다양성 분 석에 대한 연구결과만 보고되고 있다(Cho et al. 2009). 그러 나 AFLP 마커 보다 반복 재현성이 높은 microsatellite 마커를 활용하여 국내에 자생하고 있는 돌배나무를 대상으로 유전적 다양성 분석에 활용된 연구 결과는 거의 없는 실정이며, 이러 한 연구결과는 국내 배 품종 육성에 돌배나무 유전자원의 활 용 가능성에 대한 기초 자료로 매우 유용하게 사용되리라 사 료된다.
따라서 본 연구에서는 국내 전국 각지에서 수집된 돌배나 무의 형태적 특성 조사 및 microsatellite 마커를 이용한 유전 적 다양성 분석 등에 대한 일련의 연구를 수행하여 얻어진 결 과를 보고하는 바이다.
분석재료 및 형태적 특성 조사
본 연구에서는 국립산림과학원 수집, 보존되고 있는 시험 림에서 돌배나무 62점을 공시하였다(Table 1). 공시 수집 개 체 중에서 특성 조사가 가능하였던 24점에 대하여 수집 개체 당 5반복으로 하여 6개의 화기 관련 특성(과지당 꽃 수, 자방 과 수술의 길이, 악통 길이와 직경, 화서 길이), 3개의 잎 특성 (잎의 길이와 폭 및 엽병 길이), 5개의 과실 특성(과실 길이와 폭, 과병 길이, 과실무게, 당도)을 조사하였다.
Table 1.
Pear accessions assayed for genetic characterization using microsatellite markers.
Table 1.
No. Accession name Collection site

1 Chiak 1 Gangwon-do Wonju
2 Chiak 2 Gangwon-do Wonju
3 Chiak 3 Gangwon-do Wonju
4 Chiak 4 Gangwon-do Wonju
5 Chiak 5 Gangwon-do Wonju
6 Chiak 6 Gangwon-do Wonju
7 Dongmyeon 1 Gangwon-do Jeongseon
8 Dongmyeon 2 Gangwon-do Jeongseon
9 Nammyeon 1 Gangwon-do Jeongseon
10 Nammyeon 2 Gangwon-do Jeongseon
11 Nammyeon 3 Gangwon-do Jeongseon
12 Nammyeon 4 Gangwon-do Jeongseon
13 Imgye 1 Gangwon-do Jeongseon
14 Imgye 2 Gangwon-do Jeongseon
15 Imgye 3 Gangwon-do Jeongseon
16 Imgye 4 Gangwon-do Jeongseon
17 Imgye 5 Gangwon-do Jeongseon
18 ImgyeN1 Gangwon-do Jeongseon
19 ImgyeN2 Gangwon-do Jeongseon
20 Soksa 1 Gangwon-do Pyeongchang
21 Soksa 2 Gangwon-do Pyeongchang
22 Soksa 3 Gangwon-do Pyeongchang
23 Pyeongchang 1 Gangwon-do Pyeongchang
24 Pyeongchang 2 Gangwon-do Pyeongchang
25 Hoengseong 1 Gangwon-do Hoengseong
26 Hoengseong 2 Gangwon-do Hoengseong
27 Hoengseong 3 Gangwon-do Hoengseong
28 Gangreung Gangwon-do Gangreung
29 Taebaek Gangwon-do Taebaek
30 Hongcheon Gangwon-do Taebaek
31 Bongwha 1 Gyeongsangbuk-do Bongwha
32 Bongwha 2 Gyeongsangbuk-do Bongwha
33 Gimcheon Gyeongsangbuk-do Gimcheon
34 Mungyeong Gyeongsangbuk-do Mungyeonng
35 Uljin Gyeongsangbuk-do Uljin
36 Yeongyang Gyeongsangbuk-do Yeongyang
37 Jinsangeochi 1 Jeollanam-do Gwangyang
38 Jinsangeochi 2 Jeollanam-do Gwangyang
39 Jinsangeochi 3 Jeollanam-do Gwangyang
40 Jinsangeochi 4 Jeollanam-do Gwangyang
41 Jinsangeochi 5 Jeollanam-do Gwangyang
42 Okgok 1 Jeollanam-do Gwangyang
43 Okgok 2 Jeollanam-do Gwangyang
44 Okgok 3 Jeollanam-do Gwangyang
45 Gokseong 1 Jeollanam-do Gokseong
46 Gokseong 2 Jeollanam-do Gokseong
47 Gokseong 3 Jeollanam-do Gokseong
48 Piagol 1 Jeollanam-do Gurye
49 Piagol 2 Jeollanam-do Gurye
50 Piagol 3 Jeollanam-do Gurye
51 Wando Jeollanam-do Wando
52 Okgok-supyeong Jeollanam-do Gwangyang
53 Okryong Jeollanam-do Gwangyang
54 Muju Jeollabuk-do Muju
55 Baekryeonsa Jeollabuk-do Muju
56 Cheongsil Jeollabuk-do Jinan
57 Namyangju 1 Gyeonggi-do Namyangju
58 Namyangju 2 Gyeonggi-do Namyangju
59 Gwanggyo Gyeonggi-do Suwon
60 Pocheon Gyeonggi-do Pocheon
61 Cheongseori Rural development administration
62 Yeongmokri Rural development administration
Microsatellite 분석
공시 수집종의 잎 조직을 액체질소를 이용하여 마쇄한 다음 NucleoSpin®PlantII (Macherey-Nagel Cat. 740 770.250, Germany) 키트를 이용하여 게놈 DNA를 분리 하였다. 추출 된 DNA는 1.5% 아가로스젤에서 전기 영동하여 DNA 농도 를 확인한 후 μL당 30 ng의 농도로 희석하여 PCR 분석에 이 용하였다. 돌배나무의 특성 평가에 활용된 microsatellite 마 커는 염색체상 위치, 반복염기서열 정도 등을 고려하여 50개 를 선정하여(Bao et al. 2007, Kimura et al. 2002, Nishitani et al. 2009, Sehic et al. 2012, Wolko et al. 2010, Yamamoto et al. 2002) 프라이머의 정방향에 FAM, VIC, NED, PET (Applied Biosystem, USA)으로 형광 표지한 다음 이를 PCR 에 이용하였다. PCR 반응은 게놈 DNA 30 ng, 0.1 μM의 SSR primer, 2.0 μl dNTP mixture (2.5 mM), Taq polymerase 1.0 U, 2.5 μl의 10×PCR buffer (50 mM KCl, 20 mM Tris-HCl, pH 8.0, 2.0 mM MgCl2)에 초순수를 첨가하여 전 체 부피를 30 μl로 조절하였다. PCR (C1000, BioRad, USA) 증폭은 94°C에서 30초간 denature 한 후, 55°C에서 30초간 annealing, 72°C에서 45초간 extension을 40cycle 수행하였다. PCR 증폭 산물 1.0 μl와 탈이온된 포름아마이드(deionized formamide) 10 μl, size marker (LIZ500 size standard) 0.25 μl를 혼합하고 94°C에서 2분간 변성시킨 다음 자동염기서열 분석기(Genetic Analyzer 3130XL, Applied Biosystem, USA) 를 활용하여 전기 영동 하였다. Microsatellite 마커에 의해 증폭 된 대립 유전자의 크기는 GeneMapper (version 3.7) 컴퓨터 프 로그램(Applied Biosystem, USA)을 이용하여 분석하였다.
유전적 다양성 분석
Microsatellite 마커의 다형성 정도를 조사하기 위하여 대 립 유전자의 수, 대립 유전자의 출현빈도, 기대된 이형접합성, 관찰된 이형접합성 및 Polymorphism information content (PIC) 값은 PowerMarker (version 3.25) (Liu & Muse 2005) 를 이용하여 산출하였다. NTSYSpc (version 2.10b) (Rohlf 2000) 컴퓨터 프로그램을 이용하여 Jaccard 방법에 준하여 유전적 유사도 값을 계산한 다음 unweighted pair-group method with arithmetical average (UPGMA) (Sneath & Sokal 1973) 방 법으로 집괴 분석하고 계통도를 작성한 다음 수집종별 유전 적 다양성 및 유연 관계를 비교 분석하였다.
형태적 특성 조사
국내에서 수집된 돌배나무 유전자원 62점 중 형태적 특성 조사가 가능하였던 24점에 대하여 화기, 잎, 과실에 대한 14 가지의 형태적 특성을 조사한 바(Table 2), 자방의 길이는 ‘치 악 4’가 6.27 mm로 가장 짧았고 ‘임계 3’이 11.40 mm로 가 장 길었으며, 수술 길이의 경우 수집 종에 따라 4.82~11.24 mm까지 다양한 변이를 나타냈다. 잎의 길이는 ‘동면 1’과 ‘횡성 1’이 99 mm로 짧은 양상을 보였으며, 나머지 수집개체 는 100.45~158.07 mm까지 큰 차이를 나타냈다. 돌배나무 의 조사형질 중에서 가장 중요한 것으로 판단되는 과실의 무 게는 수집 개체에 따라서 최소 13 g에서 254 g까지 큰 변이 양상을 나타냈다. 이상의 결과를 종합해 볼 때 돌배나무의 형 태적 특성은 수집된 지역에 따라 다양한 형태적 특성 차이를 나타내는 것으로 분석되었으나, 본 연구에서 조사된 모든 형 질이 환경 조건에 따라 다양한 변이를 나타내는 양적 형질이 기 때문에 이를 이용하여 돌배 수집 종의 유전적 유연관계를 판단하기 어려울 것으로 사료되었다. Rugienius et al. (2013) 에 따르면 자연상태에서 자생하고 있는 배나무는 높은 이형 접합성을 가지고 있어 형태적 특성 조사를 통한 다양성 평가 는 정확하지 않기 때문에 분자표지에 의해 이를 해결해야 한 다고 지적한 바 있다.
Table 2.
Morphological characteristics of 24 pear accessions collected in Korea.
Table 2.
Microsatellite 분석
돌배나무 수집개체의 유전적 다양성을 분석하기 위하여 일 본(Kimura et al. 2002, Yamamoto et al. 2002), 중국(Bao et al. 2007), 폴란드(Wolko et al. 2010), 스웨덴(Sehic et al. 2012)에서 동양배나무, 서양배나무의 유전자원의 특성 평가 에 활용된 microsatellite 마커와 일본에서 동양배나무 EST에 서 개발된 신규 microsatellite 마커(Nishitani et al. 2009)의 반복염기 서열 및 다형성 정도를 고려하여 최종 50개의 프라이 머를 선정하였다. 이들 프라이머와 돌배나무 62점의 genomic DNA와 PCR하고 증폭산물을 자동염기서열분석기를 이용하 여 전기영동 하였다. 50개의 프라이머 중에서 34개는 대립유 전자의 수가 6개 이하로 적을 뿐만 아니라 PIC 값도 0.3 이하 로 낮게 나타나 돌배나무 수집종의 유전적 다양성 분석에 제 외하였다. 최종 선정된 16개의 microsatellite 프라이머와 돌배 나무 62개 수집개체에 대한 다형성 정도를 조사한 바(Table 3), microsatellite 프라이머에 따른 대립유전자의 수는 10~ 27개로 다양하게 나타났으며, 총 278개의 대립유전자가 검출 되었고, 마커 당 평균 대립유전자의 수는 17.4개로 나타났다. 한편, 유전자원의 다양성 정도를 나타내는 관찰된 이형접합성 과 기대된 이형접합성의 평균 값이 각각 0.776과 0.850으로 높게 나타났고, 공시 수집 개체에 대한 각 마커 별 유전적 다 형성 정도를 나타내는 PIC 값도 0.658~0.893까지 높은 범 위에 분포하였으며, 평균값도 0.836으로 아주 높은 경향을 보 였다. 그리고 16개의 microsatellite 프라이머 가운데 13개가 0.80 이상의 높은 PIC 값을 나타내어 본 연구에서 활용된 microsatellite 마커는 돌배나무 수집 개체의 특성 평가에 충 분히 활용될 수 있을 것으로 나타났다.
Table 3.
Genetic parameters detected by 16 microsatellite markers in 62 pear accessions.
Table 3.
Microsatellite designation Primers sequence (5' → 3') Repeat motif Product size (bp) No. of putative alleles Expected heterozygo sity Observed heterozygo sity PIC value

BGA35 F: PET-AGAGGGAGAAAGGCGATT (AG)8 121-145 12 0.696 0.387 0.667
R: GCTTCATCACCGTCTGCT
BGT23b F: FAM-CACATTCAAAGATTAAGAT (TC)18.5 180-248 21 0.899 0.790 0.891
R: ACTCAGCCTTTTTTTCCCAC
NB103a F: FAM-CACATTCAAAGATTAAGAT (AG)31 75-129 20 0.878 0.758 0.868
R: ACTCAGCCTTTTTTTCCCAC
NH001c F: NED-AATACTAATCCTTTTTGCTAA (GA)21 106-144 16 0.868 0.774 0.854
R: TCCATTCAATCTGTCTCGGTC
NH004a F: FAM-AGGATGGGACGAGTTTAGAG (GA)19 76-119 13 0.764 0.710 0.735
R: CCACATCTCTCAACCTACCA
NH005b F: VIC-TGAGAAGAATTAGCCATGATGA (GA)20 316-361 22 0.893 0.855 0.885
R: TTACTACTTGCGTGCGTTCC
NH007b F: NED-TACCTTGATGGGAACTGAAC (AG)25 112-149 16 0.891 0.855 0.881
R: AATAGTAGATTGCAATTACTC
NH009b F: PET-CCGAGCACTACCATTGA (AG)20 134-161 14 0.857 0.839 0.841
R: CGTCTGTTTACCGCTTCT
ENH036 F: FAM-TCCCTGCCAAACATTTTCTCTAC (CT)17 128-168 10 0.844 0.867 0.827
R: TTCCTGCCCAAGAGGTATCATAA
NH014a F: VIC-CAAACCTAACCCTAAATACC (GA)17 55-102 17 0.894 0.742 0.885
R: TGTTCATATATTCATCACTC
NH015a F: NED-TTGTGCCCTTTTTCCTACC (AG)19 100-144 18 0.879 0.823 0.867
R: CTTTGATGTTACCCCTTGCTG
ENH009 F: PET-CCTCAGCCTTGATCATGCAC (CT)11.5C 135-200 26 0.894 0.968 0.887
R: GGAAGTTGTGCTAGCAAAGAAGC (CT)10
ENH026 F: FAM-ACTCCCAATCTGCCTTACAAACA (CT)16.5 136-176 14 0.869 0.823 0.856
R: GCGTTGAGTGACCTCTTTCATTT
ENH029 F: VIC-GGAAGTTGTGCTAGCAAAGAAGC (CT)23 169-232 27 0.894 0.807 0.887
R: GCCTGTTTCCACTTATGCTCACT
ENH032 F: NED-AACTGGAGGGAGCTAAGGAAATG (CT)17 118-144 10 0.677 0.823 0.658
R: CATCAAACAAAACTAGCCGAACC
ENH033 F: PET-CCTGAGGTTATTGACCCAAAAGA (TC)15.5 151-211 22 0.899 0.742 0.893
R: GGTGGATACTCACTCAGTTGGAAA

Total 278 13.596 12.419 13.382

Mean 17.375 0.850 0.776 0.836
Microsatellite 마커에 따른 각 대립유전자의 분포 빈도를 조사한 바(Fig. 1), ENH009는 192 bp에서 전체 검출된 대립 유전자 124개중 23.4%인 29개가 검출되었고, ENH033은 161 bp에서 전체 대립유전자의 24.2%가 분포되는 것으로 분 석되었으며, 나머지 12개 마커(BGT23b, NB103a, NH001c, NH004a, NH005b, NH007b, NH009b, ENH036, NH014a, NH015a, ENH026, ENH029)의 대립유전자의 분포 빈도도 ENH009와 ENH033과 같이 고르게 분포하는 양상을 나타내 었다. 그러나 BGA35와 ENH032의 경우 대립 유전자의 수가 10개 이상이더라도 특정한 위치에서 대립유전자가 50%이상 분포하여 PIC 값이 다른 마커에 비해 낮은 경향을 보였다.
Fig. 1.
Histogram depicting the alleles at 2 microsatellite loci, ENH009 (A) and ENH033 (B). Base-pair size (abscissa) and the frequency at each allele (ordinate) detected by 62 pear accessions.
KJBS-47-45_F1.gif
Microsatellite 마커를 이용한 배나무 유전자원의 다형성 평가시 대립유전자의 수, 관찰된 이형접합성, 기대된 이형접 합성 및 PIC 값으로 유전적 변이 양상을 비교할 수 있다. 일 본의 Kimura et al. (2002)이 서양계와 동양계 배나무 등 다 양한 품종을 활용하여 9개의 SSR 마커로 분석하였을 때 평균 대립유전자의 수는 14.8개임을 보고한 이래, 중국의 Bao et al. (2007)은 6개의 SSR 마커로 중국, 일본, 한국 등에서 수 집한 98개의 배나무 유전자원 및 품종을 검정한 결과 평균 28 개의 대립 유전자가 검출됨을 보고하였다. 폴란드의 Wolko et al. (2010)도 6개의 유럽형 배나무와 2개의 야생배나무를 19개의 SSR 마커로 분석시 5~11개의 대립유전자가 분석됨 을 제시하였다. 본 연구에서도 국내에서 자생하고 있는 돌배 나무 수집개체를 16개의 microsatellite 마커로 분석시 대립유 전자의 수가 10~27개까지 다양하게 분포하며 평균 대립유 전자의 수도 17.4개로 높게 나타나 일본, 중국, 폴란드 등의 연구자에 의해 수행한 연구결과와 유사한 microsatellite 변이 양상을 나타내었다. 특정 마커에서 서로 다른 대립유전자를 가지는 관찰된 이형접합성 값은 0.776으로 일본과 중국에서 분석한 결과 보다 다소 높은 값을 나타내었는데, 이러한 연구 결과는 우리나라에서 자생하는 돌배나무의 경우 번식 과정에 서 자가불화합성 관련 유전자의 활성 여부와 실생묘 상태의 번식 등과 같은 다양한 요인에 의해 유전적 변이가 발생했기 때문에 나타난 결과라고 사료된다. 한편 본 연구에서 돌배나 무 수집개체의 기대된 이형접합성 값이 관찰된 이형접합성 값보다 다소 높게 나타났는데, 이는 돌배나무가 자가불화합성 에 의해 타가수정을 주로 하지만 극히 낮은 빈도로 자가수정 이 일어나기 때문에 나타난 현상이며 이와 유사한 연구결과 를 Bao et al. (2007)Miranda et al. (2010)Rugienius et al. (2013)도 이와 유사한 연구 결과를 보고한 바 있다. 그 리고 본 연구에서 16개의 microsatellite 마커는 분석된 평균 PIC 값은 0.836으로 아주 높게 나타나 돌배나무의 유전적 다 양성 분석뿐만 아니라 우리나라 배나무 품종의 지문화 작업 에도 매우 유용하게 활용될 수 있을 것으로 사료되었다.
유전적 유연관계 분석
16개의 microsatellite 마커를 이용하여 278개 대립유전자 의 크기를 기준으로 돌배나무 수집 개체에 대한 계통도를 작 성한 바(Fig. 2), 공시 수집 개체의 전체 유사도 지수는 0.09~ 1.00의 범위에 속하였고, microsatellite 마커의 유전자형에 의해 2개의 대그룹으로 구분되었다. Ⅰ그룹은 강원도에서 수 집된 ‘치악1’ 외 12점, 경북의 ‘봉화1,2’, ‘김천’과 ‘문경’에서 수집된 4점, 전남에서 수집된 ‘진상어치1’외 12점, 전북의 ‘무주’와 ‘진안’에서 수집된 3점, 경기도의 ‘남양주1,2’ 및 광 교산에서 수집된 ‘광교’ 등 3점과 농촌진흥청 원예특작과학원 에서 수집 보존된 ‘청서리’가 속하였으며, 유사도 지수 0.17 에서 3개의 소그룹으로 구분되었다. Ⅰ-1 그룹에 속하는 21 개의 수집 개체 중에서 전남 지역에서 수집된 11점, 강원도에 서 수집된 5점, 경북 지역에서 수집된 2점, 경기도에서 수집 된 2점, 농촌진흥청의 ‘청서리’가 분포하였으며, 특히 전남 광 양 지방에서 수집된 ‘진상어치2’, ‘진상어치5’, ‘옥곡3’, ‘옥곡 수평’, ‘옥룡’ 은 microsatellite 마커의 유전자형에 의해 구분 이 되지 않았다. I-2 그룹은 강원도에서 수집된 ‘치악2’외 7 점, 경북 봉화 지역에서 수집된 ‘봉화1’, 전남 광양 지방에서 수집된 ‘옥곡2’와 구례에서 수집된 ‘피아골3’, 전북 무주에서 수집된 ‘무주’와 ‘백련사’, 진안 지역에서 수집된 ‘청실’, 경기 도 수원 광교산에서 수집한 1점이 분포하였다. 강원도에서 수 집된 ‘치악2’와 ‘속사2’의 경우 100%의 유전적 유사도를 나 타냈으며, I-3 그룹은 ‘봉화2’ 하나의 수집종이 분포하였다.
Fig. 2.
Phenogram of 62 pear accessions resulting from UPGMA analysis based on Jaccard’s similarity coefficient. The numbers (1 to 62) at right side refer to the list of pear accessions in Table 1.
KJBS-47-45_F2.gif
II그룹은 강원도에서 수집된 ‘치악3’외 16점, 경북 울진과 영양에서 수집된 2점, 전남 광양 지방에서 수집된 ‘진상어치 3’외 3점과 경기도 포천에서 수집된 1점과 농촌진흥청에서 보존되고 있는 ‘영목리’ 가 분포하였으며, 모든 돌배나무 유 전자원이 microsatellite 마커의 유전자형에 의해 뚜렷하게 구 분되었다. II그룹은 유전적 유사도 지수 0.13에서 2개의 소그 룹으로 구분할 수 있었으며, II-1 그룹은 ‘치악3’, ‘임계N1’, ‘임계1’, ‘속사1’, ‘진상어치3’, ‘곡성3’, ‘포천’ 이 분포하였 고, II-2그룹은 강원도에서 수집된 13점, 경북 지방에서 수집 된 2점, 전남 지역에서 수집된 2점, 농촌진흥청의 ‘영목리’가 포함되는 양상을 나타냈다. 이상의 결과를 분석해볼 때, 강원 도에서 수집된 종이 I, II 그룹 내에 고르게 분포하는 양상 을 나타내어 다른 지역에서 수집된 종보다 유전적 다양성이 높은 것으로 분석되었다.
Microsatellite 마커를 이용한 배나무 유전자의 특성 평가 는 일본에서 6 Pyrus 속 60품종에 대하여 9개의 SSR 마커에 서 검출된 133개 대립유전자를 활용하여 군집 분석하였을 때 유럽형, 중국형, 일본형 품종군으로 그룹화 됨을 보고한 이래 (Kimura et a1. 2002), 중국에서 6개의 SSR marker를 98개 배나무 유전자원 및 품종을 10개의 대그룹으로 구분할 수 있 음을 제시하였다(Bao et al. 2007). 스페인의 경우 141개의 배나무 유전자원 평가에 8개의 SSR 마커에서 검출된 97개의 대립유전자를 이용하여 군집분석을 실시하였을 때 3개의 대 그룹으로 구분할 수 있음을 보고하였다. 본 연구에서도 우리 나라에서 자생하고 있는 돌배나무 수집개체를 대상으로 16개 의 microsatellite 마커에서 검출된 278개의 대립유전자를 활 용하였을 때 2개의 대그룹으로 그룹화할 수 있었다. 특히, 강 원도에서 수집된 개체는 전체 그룹 내에 고르게 분포하는 양 상을 보였으며, 전남지방에서 수집한 개체의 경우 17개 수집 개체 중에서 ‘진상어치3’, ‘곡성3’, ‘피아골1’, 피아골2’를 제 외한 13개 수집종이 I그룹 내에 분포하는 것으로 나타나 수 집된 지역에 따라 유전적 변이 정도가 다른 것으로 분석되었 다. 이러한 결과는 돌배나무 수집 지역의 지리적 여건, 기상환 경에 따른 자가불합화성의 타파 여부, 실생묘 번식에 따른 유 전적 다양성 발생 등과 같은 여러 가지 요인이 관여하였기 때 문에 나타난 결과라고 사료된다. 그리고 일부 유전자원의 경 우 278개의 대립유전자를 활용해도 구분이 되지 않았는데 이 에 대해서 분석 마커의 수를 확대하거나 형태적 특성 검정을 병행하여 수집개체의 동일성 여부를 검토해야 될 것으로 사료 된다. 한편, 본 연구에서 돌배나무 수집개체에 활용된 microsatellite 마커 16개는 대립유전자의 수도 많을 뿐만 아니라 PIC 값도 높게 나타나 이를 우리나라 배나무 품종 식별에 활 용가능성 여부를 탐색한다면 실용적 활용 가치는 한층 확대 될 것으로 판단된다.
이 연구의 목적은 형태적 특성과 microsatellite 마커를 이 용하여 국내에서 자생하고 있는 돌배나무 유전자원의 유전적 다양성 평가를 위하여 수행하였다. 돌배나무 14개의 형태적 특성을 조사한 바 수집종간에 높은 변이성을 나타냈으나 환 경의 영향을 많이 받는 양적형질이기 때문에 정확한 특성평 가가 어려웠다. 50개의 microsatellite 마커를 이용하여 62개 돌배나무 수집개체에 다형성 정도가 높은 16개를 선정하였다. 이들 마커와 돌배나무 유전자원 62점을 검정하였을 때 총 284개의 대립유전자가 나타났으며, 마커에 따라 10-27개까지 다양한 분포 양상을 나타냈다. 16개 마커의 평균 PIC 값과 관 찰된 이형접합성은 각각 0.836와 0.776로 나타났다. 돌배 62 개 수집개체를 microsatellite 마커에 의해 나타난 대립유전자 를 근거로 Jaccard 방법에 따라 산출된 유전적 유사도는 0.09 ~1.00까지 넓은 범위에 속하였고, UPGMA 방법에 따라 군 집분석을 실시하였을 때, 2개의 그룹으로 크게 나누어졌으며 55개 돌배나무 수집개체가 구분되는 것으로 나타났다. 본 연 구 결과는 돌배나무 수집개체의 유전적 다양성과 유연관계 평가를 통해 유용한 유전자원에 대한 정보를 제공하는데 유 용하게 활용될 수 있을 것이다.
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Genetic Diversity of Wild Pear Accessions Collected in Korea
Korean. J. Breed. Sci.. ;47(1):45-53.   Published online March 31, 2015
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Korean. J. Breed. Sci.. ;47(1):45-53.   Published online March 31, 2015
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Genetic Diversity of Wild Pear Accessions Collected in Korea
Image Image
Fig. 1. Histogram depicting the alleles at 2 microsatellite loci, ENH009 (A) and ENH033 (B). Base-pair size (abscissa) and the frequency at each allele (ordinate) detected by 62 pear accessions.
Fig. 2. Phenogram of 62 pear accessions resulting from UPGMA analysis based on Jaccard’s similarity coefficient. The numbers (1 to 62) at right side refer to the list of pear accessions in Table 1.
Genetic Diversity of Wild Pear Accessions Collected in Korea

Pear accessions assayed for genetic characterization using microsatellite markers.

No. Accession name Collection site

1 Chiak 1 Gangwon-do Wonju
2 Chiak 2 Gangwon-do Wonju
3 Chiak 3 Gangwon-do Wonju
4 Chiak 4 Gangwon-do Wonju
5 Chiak 5 Gangwon-do Wonju
6 Chiak 6 Gangwon-do Wonju
7 Dongmyeon 1 Gangwon-do Jeongseon
8 Dongmyeon 2 Gangwon-do Jeongseon
9 Nammyeon 1 Gangwon-do Jeongseon
10 Nammyeon 2 Gangwon-do Jeongseon
11 Nammyeon 3 Gangwon-do Jeongseon
12 Nammyeon 4 Gangwon-do Jeongseon
13 Imgye 1 Gangwon-do Jeongseon
14 Imgye 2 Gangwon-do Jeongseon
15 Imgye 3 Gangwon-do Jeongseon
16 Imgye 4 Gangwon-do Jeongseon
17 Imgye 5 Gangwon-do Jeongseon
18 ImgyeN1 Gangwon-do Jeongseon
19 ImgyeN2 Gangwon-do Jeongseon
20 Soksa 1 Gangwon-do Pyeongchang
21 Soksa 2 Gangwon-do Pyeongchang
22 Soksa 3 Gangwon-do Pyeongchang
23 Pyeongchang 1 Gangwon-do Pyeongchang
24 Pyeongchang 2 Gangwon-do Pyeongchang
25 Hoengseong 1 Gangwon-do Hoengseong
26 Hoengseong 2 Gangwon-do Hoengseong
27 Hoengseong 3 Gangwon-do Hoengseong
28 Gangreung Gangwon-do Gangreung
29 Taebaek Gangwon-do Taebaek
30 Hongcheon Gangwon-do Taebaek
31 Bongwha 1 Gyeongsangbuk-do Bongwha
32 Bongwha 2 Gyeongsangbuk-do Bongwha
33 Gimcheon Gyeongsangbuk-do Gimcheon
34 Mungyeong Gyeongsangbuk-do Mungyeonng
35 Uljin Gyeongsangbuk-do Uljin
36 Yeongyang Gyeongsangbuk-do Yeongyang
37 Jinsangeochi 1 Jeollanam-do Gwangyang
38 Jinsangeochi 2 Jeollanam-do Gwangyang
39 Jinsangeochi 3 Jeollanam-do Gwangyang
40 Jinsangeochi 4 Jeollanam-do Gwangyang
41 Jinsangeochi 5 Jeollanam-do Gwangyang
42 Okgok 1 Jeollanam-do Gwangyang
43 Okgok 2 Jeollanam-do Gwangyang
44 Okgok 3 Jeollanam-do Gwangyang
45 Gokseong 1 Jeollanam-do Gokseong
46 Gokseong 2 Jeollanam-do Gokseong
47 Gokseong 3 Jeollanam-do Gokseong
48 Piagol 1 Jeollanam-do Gurye
49 Piagol 2 Jeollanam-do Gurye
50 Piagol 3 Jeollanam-do Gurye
51 Wando Jeollanam-do Wando
52 Okgok-supyeong Jeollanam-do Gwangyang
53 Okryong Jeollanam-do Gwangyang
54 Muju Jeollabuk-do Muju
55 Baekryeonsa Jeollabuk-do Muju
56 Cheongsil Jeollabuk-do Jinan
57 Namyangju 1 Gyeonggi-do Namyangju
58 Namyangju 2 Gyeonggi-do Namyangju
59 Gwanggyo Gyeonggi-do Suwon
60 Pocheon Gyeonggi-do Pocheon
61 Cheongseori Rural development administration
62 Yeongmokri Rural development administration

Morphological characteristics of 24 pear accessions collected in Korea.

Genetic parameters detected by 16 microsatellite markers in 62 pear accessions.

Microsatellite designation Primers sequence (5' → 3') Repeat motif Product size (bp) No. of putative alleles Expected heterozygo sity Observed heterozygo sity PIC value

BGA35 F: PET-AGAGGGAGAAAGGCGATT (AG)8 121-145 12 0.696 0.387 0.667
R: GCTTCATCACCGTCTGCT
BGT23b F: FAM-CACATTCAAAGATTAAGAT (TC)18.5 180-248 21 0.899 0.790 0.891
R: ACTCAGCCTTTTTTTCCCAC
NB103a F: FAM-CACATTCAAAGATTAAGAT (AG)31 75-129 20 0.878 0.758 0.868
R: ACTCAGCCTTTTTTTCCCAC
NH001c F: NED-AATACTAATCCTTTTTGCTAA (GA)21 106-144 16 0.868 0.774 0.854
R: TCCATTCAATCTGTCTCGGTC
NH004a F: FAM-AGGATGGGACGAGTTTAGAG (GA)19 76-119 13 0.764 0.710 0.735
R: CCACATCTCTCAACCTACCA
NH005b F: VIC-TGAGAAGAATTAGCCATGATGA (GA)20 316-361 22 0.893 0.855 0.885
R: TTACTACTTGCGTGCGTTCC
NH007b F: NED-TACCTTGATGGGAACTGAAC (AG)25 112-149 16 0.891 0.855 0.881
R: AATAGTAGATTGCAATTACTC
NH009b F: PET-CCGAGCACTACCATTGA (AG)20 134-161 14 0.857 0.839 0.841
R: CGTCTGTTTACCGCTTCT
ENH036 F: FAM-TCCCTGCCAAACATTTTCTCTAC (CT)17 128-168 10 0.844 0.867 0.827
R: TTCCTGCCCAAGAGGTATCATAA
NH014a F: VIC-CAAACCTAACCCTAAATACC (GA)17 55-102 17 0.894 0.742 0.885
R: TGTTCATATATTCATCACTC
NH015a F: NED-TTGTGCCCTTTTTCCTACC (AG)19 100-144 18 0.879 0.823 0.867
R: CTTTGATGTTACCCCTTGCTG
ENH009 F: PET-CCTCAGCCTTGATCATGCAC (CT)11.5C 135-200 26 0.894 0.968 0.887
R: GGAAGTTGTGCTAGCAAAGAAGC (CT)10
ENH026 F: FAM-ACTCCCAATCTGCCTTACAAACA (CT)16.5 136-176 14 0.869 0.823 0.856
R: GCGTTGAGTGACCTCTTTCATTT
ENH029 F: VIC-GGAAGTTGTGCTAGCAAAGAAGC (CT)23 169-232 27 0.894 0.807 0.887
R: GCCTGTTTCCACTTATGCTCACT
ENH032 F: NED-AACTGGAGGGAGCTAAGGAAATG (CT)17 118-144 10 0.677 0.823 0.658
R: CATCAAACAAAACTAGCCGAACC
ENH033 F: PET-CCTGAGGTTATTGACCCAAAAGA (TC)15.5 151-211 22 0.899 0.742 0.893
R: GGTGGATACTCACTCAGTTGGAAA

Total 278 13.596 12.419 13.382

Mean 17.375 0.850 0.776 0.836
Table 1. Pear accessions assayed for genetic characterization using microsatellite markers.
Table 2. Morphological characteristics of 24 pear accessions collected in Korea.
Table 3. Genetic parameters detected by 16 microsatellite markers in 62 pear accessions.