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국내에서 수집된 국화 품종에 대한 SSR 분자표지 Database 구축

Construction of SSR Marker Database of Chrysanthemum Varieties Collected in Korea

Eun-Jo Shim1, Eun-Jung Heo2, Moo-Kyoung Yoon3, Eun-Hee Soh1, Jee-Hwa Hong4,*
Korean Journal of Breeding Science 2015;47(4):366-375.
Published online: November 30, 2015

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4 국립농산물품질관리원 시험연구소 품질조사과

4 Division of Quality Examination, Experiment Research Institute, National Agricultural Products QualityManagement Service, Gimcheon 39660, Korea

*Corresponding author (hongjh19@korea.kr, Tel: +82-54-912-0230, Fax: +82-54-912-0227)
• Received: September 1, 2015   • Revised: September 24, 2015   • Accepted: September 25, 2015

© Korean Society of Breeding Science All rights reserved

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  • Chrysanthemum (Dendranthema grandiflourm Kitamura) is a member of the Asteraceae and one of the important horticultural crops. The objective of this study was to construct a DNA profile database for identification of chrysanthemum varieties using simple sequence repeat (SSR) markers. In order to select SSR markers for the variety identification, we screened 587 SSR primers using 20 varieties. Among them, 27 SSR markers showed polymorphism. We finally selected 14 SSR markers showing peak clearance, high polymorphism and reproducibility in 20 varieties. In conclusion, DNA profile database for 147 chrysanthemum varieties were constructed by 14 SSR markers. A total of 79 SSR alleles were detected and three to ten alleles were detected with an average of 5.6 alleles per locus. The polymorphism information content value ranged 0.287 ~ 0.785 with an average of 0.598. Genetic relationship revealed that genetic distance of 147 varieties ranged from 0.44 to 1.00. The 143 varieties among 147 varieties were distinguished by 14 SSR markers but the 2 varieties developed by mutation breeding and natural variation were not distinguished from original varieties. These constructed SSR profile database will be useful for the selection of similar varieties for candidate variety and for solving problem relating to seed dispute and infringement of plant breeder’s right.
국화(Dendranthema grandiflourm Kitamura)는 국화과 (Asteraceae)에 속하고 육종목표에 따라 절화, 화분국, 화단국 등 다양한 형태로 육성되어 재배되고 있다(Anderson 2007). 국내에서 국화는 다양한 화색과 화형을 갖춘 품종들이 개발 되어 2015년 6월 30일 현재 국화의 품종보호 등록 품종수는 638품종(http://www.seed.go.kr)으로 화훼작물 중 장미 다음 으로 보호 등록된 품종수가 많은 작물이다. 따라서 품종보호 침해 등과 같은 지식재산권 분쟁이 발생할 확률이 높고, 형태 적으로 유사한 품종간의 구별성 확인 등을 통한 육종가의 품 종보호권 보호가 절실히 요구된다. 국화 품종을 효율적으로 관리하고, 식별하기 위한 방법 중의 하나가 품종식별 분자표 지를 이용하여 이를 데이터베이스화하는 것이며 국제식물신 품종보호연맹(UPOV, international union for the protection of new varieties of plants)에서는 분자표지 선발과 이를 이 용한 DNA profile 데이터베이스 구축에 관한 유전자 분석 지 침서를 제정한바 있다(UPOV 2010). 국립종자원에서는 UPOV 에서 제안한 유전자 분석 지침서를 기반으로 고추(Kwon et al. 2013), 멜론(Kwon & Hong 2014), 상추(Hong et al. 2013a) 등의 27개 작물에 대하여 게놈 내 단순 반복되는 염 기의 변이를 이용하는 simple sequence repeat (SSR) 분자표 지를 이용하여 DNA profile 데이터베이스를 구축하여 품종 보호 출원품종의 대조품종 선정과 유통종자 및 분쟁종자의 품종진위 검정 등에 활용하고 있는 실정이다.
국화의 품종식별에 관한 국내외 연구동향으로 중국에서는 18품종을 대상으로 randomly amplified polymorphic DNA (RAPD) 분자표지를 이용하여 유연관계를 분석하였고(Helan et al. 2002), amplified fragment length polymorphism (AFLP) 분자표지를 이용하여 45개 품종에 대한 유전적 다양성 연구 결과를 보고하였다(Jie et al. 2007). 최근에는 next generation sequencing(NGS) 분석법을 통해 개발한 expressed sequence tag(EST) 유래 SSR 분자표지를 이용하여 종(species) 식별에 대한 연구결과가 보고되었다(Wang et al. 2013). 국내에서는 SSR 마커를 이용하여 85품종에 대한 식별 연구가 수행되었 고(Hong et al. 2013b), EST-SSR 분자표지를 이용하여 50품 종에 대한 식별 연구 결과가 보고된 바 있다(Jo et al. 2015). 국화는 경제적으로 중요한 작물이기에 분자표지 연구가 최근 많이 이루어지고 있지만 많은 품종을 대상으로 한 정밀도 높 은 데이터베이스 구축은 미비한 실정이기 때문에 품종보호제 도, 종자분쟁 해결 등에 분자표지를 실용적으로 활용하기 위 해서는 DNA profile 데이터베이스 구축을 통한 체계적인 품 종식별 체계 확립이 필요하다고 판단되었다.
따라서 본 연구에서는 다양한 분자표지 분석방법들 중 SSR 분자표지를 활용하여 국내에서 수집된 147개 국화의 품 종에 대한 품종식별 분자표지 선발과 품종별 DNA profile 데 이터베이스 구축을 실시하였다.
공시품종 및 genomic DNA 분리
공시품종은 국화 147개 품종을 충청남도농업기술원 화훼연구 소와 국립종자원 서부지원에서 수집하여 DNA profile 데이터베 이스 구축에 활용하였다(Table 1). 공시품종의 DNA 추출은 본 엽 3 ~ 4매를 채취하여 Nucleospin®PlantII(Macherey-Nagel Cat. 740770.250) Kit를 이용하여 분리하였다. 추출된 DNA 는 1.5% agarose gel에서 전기영동 하여 DNA 농도를 확인한 후 μl 당 20 ng의 농도로 정량하여 PCR 분석에 이용하였다.
Table 1.
Chrysanthemum varieties used for genetic relationship analysis using simple sequence repeat markers.
Table 1.
No. Variety name Type classification No. Variety name Type classification
1 Borami Spray 45 Delidantepurple Spray
2 Mujigae Spray 46 Delibiscari Spray
3 Hwiparam Spray 47 Delitents Spray
4 Yesmorning Spray 48 Deliaragon Spray
5 Magic Spray 49 Deliquebec Spray
6 Yellowcap Spray 50 Delimarimo Spray
7 Ilweol Spray 51 Delithm Spray
8 Cherryblossom Spray 52 Lovemine Spray
9 Forestaroma Spray 53 Yeosim Spray
10 Secretpink Spray 54 Magicstar Spray
11 OrangeND Spray 55 Delmonte Spray
12 PeachND Spray 56 Moulinrouge Spray
13 BrightND Spray 57 SunnyND Spray
14 PowerND Spray 58 BounceND Spray
15 Dreamround Spray 59 Athos(Sp.) Spray
16 Firepink Spray 60 HappyND Spray
17 GraceND Spray 61 Boramiorange Spray
18 Kingfisher Spray 62 Azma Spray
19 Yesruby Spray 63 Sasineyellow Spray
20 Vogue Spray 64 Nostar Spray
21 CutieND Spray 65 Sasinecream Spray
22 Merrybell Spray 66 Caesarcream Spray
23 Dreamhappy Spray 67 Seireine Spray
24 Dreamriver Spray 68 Seiribbu Spray
25 Deliacream Spray 69 Seimyra Spray
26 Purplecap Spray 70 Seited Spray
27 Lollipop Spray 71 Seiastron Spray
28 Pinkmam Spray 72 Seielza Spray
29 Argus Spray 73 Sweetbongbong Spray
30 Purplecone Spray 74 Leopard Spray
31 Biarittz P Spray 75 Sweetsnow Spray
32 Goldrich Spray 76 Ford Spray
33 Monalisa Spray 77 Sweetsun Spray
34 Goldenboy Spray 78 Castelli Spray
35 Bacardifill Spray 79 Sweetcandy Spray
36 Pinkyboy Spray 80 Artirisingsun Spray
37 Gloria Spray 81 Pinky Spray
38 Vatican Spray 82 Artipurplelady Spray
39 Yeonja Spray 83 Artidarkchocolate Spray
40 Naos Spray 84 Yesholic Spray
41 Sasine Spray 85 Jazz Spray
42 Neseteo Spray 86 Yellowdream Spray
43 Seifrillgreen Spray 87 Spybelly Spray
44 Delibarca Spray 88 Yellowmarble Spray
89 Seiagness Spray 119 Zemblalime Standard
90 Yellowkid Spray 120 Armadolis Pot mum
91 Champy Spray 121 Aamflu Pot mum
92 Orangedream Spray 122 Myruby Pot mum
93 Zarita Spray 123 Vino Pot mum
94 Orangekid Spray 124 Mygirl Pot mum
95 Hirario Spray 125 Pinkeye Pot mum
96 Zembla Spray 126 Mysoul Pot mum
97 Cutiepink Spray 127 Gayagold Pot mum
98 Cheeks Spray 128 Gayaorange Pot mum
99 Fireyellow Spray 129 Gayaglory Pot mum
100 Karisyellow Spray 130 Gayayellow Pot mum
101 Pinkycharis Spray 131 Gayaneon Pot mum
102 Yellowpangpang Spray 132 Gayaruby Pot mum
103 Pinkpangpang Spray 133 Purpleegg Pot mum
104 Rubyna Spray 134 Juliet Pot mum
105 Hongpa Spray 135 Peacecutie Pot mum
106 Hibaeksan Standard 136 Gayapink Pot mum
107 Sumi Standard 137 Mygold Pot mum
108 Baekma Standard 138 Mysunny Pot mum
109 Shinma Standard 139 Myred Pot mum
110 Geumhwa Standard 140 Gayasense Pot mum
111 Boryeonghwangkuk Standard 141 Mywing Pot mum
112 GreenND Standard 142 Mycream Pot mum
113 Athos(St.) Standard 143 Candyball Garden mum
114 Deliakari Standard 144 Dasomball Garden mum
115 Whiterunner Standard 145 Maruball Garden mum
116 Pingpong Standard 146 Madangball Garden mum
117 Seolmi Standard 147 Geumbangul Garden mum
118 Ecogreen Standard
Primer 확보 및 SSR 분석
국화 품종식별에 효과적인 SSR 분자표지를 선발하기 위해 서 국내외에서 개발된 575개 primer(Wang et al. 2013), 12 개의 primer(Khaing et al. 2013) 총 587개의 primer를 이용 하여 ‘Borami(보라미)’, ‘Mujigae(무지개)’, ‘Hwiparam(휘파 람)’, ‘Yesmorning(예스모닝)’, ‘Hibaeksan(하이백산)’, ‘Magic (매직)’, ‘Yellowcap(옐로우캡)’, ‘Sumi(수미)’, ‘Ilweol(일월)’, ‘Cherryblossom (체리블럿섬)’, ‘Forestaroma(포레스트아로마)’, ‘Secretpink(시크릿핑크)’, ‘Baekma(백마)’, ‘OrangeND(오렌 지엔디)’, ‘PeachND(피치엔디)’, ‘BrightND(브라이트엔디)’, ‘PowerND(파워엔디)’, ‘Dreamround(드림라운드)’, ‘Firepink (파이어핑크)’, ‘Shinma(신마)’ 20개 품종을 대표품종으로 선 정하여 다형성을 분석하였다. PCR 반응을 통한 증폭산물은 QIAxcel Advanced System(QIAGEN, Germany)을 이용하 여 전기영동하였고 컴퓨터 프로그램(QIAxcel ScreenGel)을 활용하여 대표품종 간에 대립유전자의 차이를 분석하여 다형 성을 보이는 분자표지를 1차 선발하였다. 선발된 primer는 정 방향에 FAM, VIC, NED, PET (Applied Biosystems, USA) 으로 형광 표지한 다음 국화 147품종(Table 1)에 대한 DNA profile 데이터베이스 구축에 이용하였다.
PCR 반응은 국화 genomic DNA 20 ng, 0.5 μM의 SSR primer, 2 μl dNTP mixture(2.5 mM), Taq polymerase 1 unit, 2.5 μl의 10 X PCR buffer(50 mM KCl, 20 mM Tris-HCl, pH 8.0, 2 mM MgCl2)에 증류수를 첨가하여 전체 부피를 25 μl로 반응액을 조성하였다. PCR(GeneAmp® PCR System 9700) 증폭은 95℃에서 5분 동안 pre-denaturation 한 후, 95℃에서 30초간 denaturation, 55℃에서 30초간 annealing, 72℃에서 45초간 extension을 40 cycle 수행하였 고, 72℃에서 10분간 final-extension을 수행하였다. PCR이 완료된 후 1.0 ~ 2.0 μl의 증폭산물을 2% agarose gel에서 전 기영동하여 증폭산물을 확인한 후, 초순수 220 μl에 PCR 산 물 1.0 μl를 희석하였다. 희석된 PCR 증폭산물 1.0 μl는 탈이 온 된 포름아마이드(deionized formamide) 10 μl, size marker(LIZ500 size standard) 0.25 μl를 혼합한 다음 95℃ 에서 2분간 denaturation 하였다. 변성된 PCR 증폭산물은 자동염 기서열분석(Genetic Analyzer 3130XL, Applied Biosystems, USA)를 활용하여 전기영동한 후 GeneMapper (ver. 3.7) 컴 퓨터 프로그램(Applied Biosystems, USA)을 이용하여 SSR 분자표지에 대한 대립유전자의 크기를 분석하였다.
다형성 지수 및 품종간 유전적 유사도 분석
SSR 분자표지의 대립유전자 유무에 따라 ‘1(present)’과 ‘0(absent)’로 코드화한 후 엑셀파일에 품종별 대립유전자형 을 크기별로 정리하여 데이터베이스를 구축하였다. SSR 분자 표지의 다형성 정도를 조사하기 위하여 Anderson et al. (1993)이 보고한 공식을 이용하여 polymorphism information contents(PIC) 값을 계산하였다. NTSYSpc(ver. 2.10b) (Rohlf 2000) 컴퓨터 프로그램의 Jaccard 방법에 따라 유전 적 유사도 값을 산출하였고, unweighted pair-group method with arithmetical average(UPGMA)(Sneath & Sokal 1973) 방법으로 집괴 분석하여 품종간 유사도를 분석하였다.
SSR 분자표지 데이터베이스 구축
국화 품종간 식별이 되는 분자표지를 선발하기 위하여 SSR primer 587개를 이용하여 ‘Borami(보라미)’ 포함 20품 종을 이용하여 다형성 검정을 수행한 결과 총 587개 중 SSR 48, 127, 330과 같이 다형성을 나타내는 27개의 분자표지를 선 발하였다(Fig. 1). 선발된 27개의 분자표지는 12개의 genomic SSR과 575개의 EST-SSR 마커로부터 선발 되었다(Table 2). 국화 genomic DNA에서 개발된 SSR 분자표지의 다형성 비 율은 EST-SSR 마커에 비해 높았으나 국화 품종식별을 위해 최종 선발된 27개 SSR 분자표지 중 25개는 EST-SSR 분자 표지로 확인되었다. 본 연구에서는 SSR 마커의 선발 비율이 낮게 나타났는데 이는 Capillary 젤 전기영동 분석시 다수의 마커에서 대립유전자가 많이 나타나 복잡한 양상을 나타내었 고 이러한 마커는 선발시 제외되었기 때문인 것으로 생각된 다. 선발된 27개의 SSR 분자표지를 이용하여 DNA sequencer 젤 전기영동 분석을 통해 다형성 정도, 피크 패턴, 반복성 등을 조사하여 EST에서 유래된 14개의 SSR 분자표 지를 국화 품종별 DNA profile 데이터베이스 구축에 활용할 마커로 최종 선발하였다(Table 3). Leigh et al. (2003)은 EST-SSR 분자표지는 genomic SSR 분자표지에 비해 밴드가 명확하여 정밀도 높은 유전자형 분석에 효과적인 것으로 보 고한 바 있는데 본 연구에서도 대립유전자 크기 판독이 용이 하였다. 14개 분자표지에 의해 분석된 대립유전자 수는 총 79 개였으며, 분자표지당 대립유전자 수는 3 ~ 10개의 분포를 나 타내었고, 평균 대립유전자 수는 5.6개였다. Hong et al. (2013b)은 4개의 genomic DNA로부터 유래된 SSR 분자표 지를 이용하여 재배국화 85품종을 분석한 결과 대립유전자 수는 11개에서 34개의 분포를 나타내었다고 보고하였다. 그 러나 본 연구에서 분석된 SSR 분자표지의 평균 대립 유전자 수는 Hong et al. (2013b)의 결과보다는 적게 나타났지만 재 배되는 국화의 배수체인 6배체(Anderson, 2007)와 유사한 수 치를 나타내었다. 그리고 Jo et al. (2015)은 16개의 EST-SSR 마커로 국화 50품종을 분석한 결과 평균 6.25개의 대립유전 자를 가진다고 보고하여 본 결과와 유사한 수치를 나타내었 다. 한편 다형성 정도를 나타내는 PIC 값은 0.287 ~ 0.785 범 위에 분포하였고, 평균 0.598를 나타내었다. Hong et al. (2013b)은 평균 PIC 값이 0.82로 높게 나타났다고 하였으나, Jo et al. (2015)는 PIC 값이 0.1516에서 0.8872의 분포로 평 균 0.603을 나타내었다고 하여 본 연구에서 나타난 PIC 값은 Jo et al. (2015)과 유사하게 나타났다. 대립유전자 수와 PIC 값의 차이는 각 연구자마다 활용된 SSR 마커의 종류가 다르 고, 공시품종간 유전적 다양성의 차이가 있기 때문에 나타난 결과로 사료된다. 국화는 배수체 작물로 대립유전자가 많아 분자표지 분석 시 마커에 따른 유전자형이 복잡한 양상을 많 이 나타내어 유전자형 분석이 어려울 수 있으나, 실제 배수체 작물인 블루베리(Hong et al. 2014a), 딸기(Hong et al. 2014b)에서도 다형성과 재현성이 높고 분석이 용이한 SSR 마커를 이용하여 품종별 DNA Profile 데이터베이스를 구축 한 사례가 있다. 배수체 작물의 경우 정밀한 유전자형 분석이 가능하다면 분자표지 기술이 품종식별과 유사품종들에 대한 품종진위 검정 등에 효과적으로 활용될 수 있을 것으로 사료 된다.
Fig. 1.
Polymorphism of 3 simple sequence repeat markers, SSR-48, SSR-127, SSR-330. The PCR products were separated using a QIAxcel Advanced System (QIAGEN). Lane 1: ‘Borami’, 2: ‘Mujigae’, 3: ‘Hwiparam’, 4: ‘Yesmorning’, 5: ‘ Hibaeksan’, 6: ‘Magic’, 7: ‘Yellowcap’, 8: ‘Sumi’, 9: ‘Ilweol’, 10: ‘Cherryblossom’, 11: ‘Forestaroma’, 12: ‘Secretpink’, 13: ‘Baekma’, 14: ‘OrangeND’, 15: ‘PeachND’, 16: ‘BrightND’, 17: ‘PowerND’, 18: ‘Dreamround’, 19: ‘Firepink’, 20: ‘Shinma’. The A, B, C indicate specific allele type.
KJBS-47-366_F1.gif
Table 2.
Simple sequence repeat markers screened for identifying polymorphism of chrysanthemum.
Table 2.
Number of screened markers Type of SSR markers Number of polymorphic markers Polymorphism (%) of amplified SSR markers SSR marker source
12 Genomic SSR of chrysanthemum 2 2/12 Khaing et al. (2013)
575 EST-SSR of chrysanthemum 25 25/575 Wang et al. (2013)
587 27 27/587 (4.6%)
Table 3.
Characteristics of the 14 simple sequence repeat markers selected for genetic characterization of chrysanthemum varieties.
Table 3.
No. Marker name Repeat motif Annealing temp. (°C) Forward sequence Reverse sequence PCR product size (bp) No. of alleles PIC value Primer labeling
1 SSR 48 (CCA)5 55 TGAGATCATTCCCAACCTCC CTAGCGTCCAAAGAATTGGC 206-214 4 0.655 VIC
2 SSR 51 (CCAA)4 55 CCCCCTCTTCTTCTTCAACC CAATAGAAAGCGCGTGACAA 197-252 8 0.785 NED
3 SSR 52 (ATA)5 55 AGTGACCCGAGCCAGATAGA CCGACAAATCATTTCCGTCT 233-247 6 0.573 FAM
4 SSR 127 (GGT)5 55 TAACAAGGGGTTTCAGCGTC TCAGGAAGAACAACCCAACC 262-273 5 0.335 FAM
5 SSR 200 (GGT)5 55 CCCAGAGAAGCGTGAGATTT TCCCCTGCTACTACCACCAC 129-154 4 0.637 VIC
6 SSR 222 (ATC)5 55 AGCTAAAACAAACAAGCGGC GCGTTAACTGTGTCGGTTGA 153-185 8 0.756 NED
7 SSR 330 (CAA)5 55 CTGTTGAGCAGTTCAGGCAC GTGTGATCGAGGCGATTTTT 204-219 6 0.597 FAM
8 SSR 332 (TGG)5 55 ACACCGAATAGGACGGACAG TTTTCTGAAGTCCCGACCAC 227-240 5 0.556 PET
9 SSR 380 (TGA)5 55 ACCAAAGGCAGCTCACAGTT CCTCCCTCACTCATCTCTGC 190-196 4 0.658 VIC
10 SSR 649 (ACC)7 55 TCTTCCTCACACGCAAACAC AGCTGCCACTCGCTATCACT 247-259 6 0.762 FAM
11 SSR 706 (AG)8 55 CGATCACCATTCTTTTCCCA CCGATAAGTTCGTCCTTGGT 208-240 7 0.613 PET
12 SSR 728 (ATG)5 55 TGGTTATGGGTGACCCTGAT AAGAAAGTGCAGGCCAAGAA 145-151 3 0.488 VIC
13 SSR 792 (TGA)6 55 AGGAAGAAGATCGACACCCA AAGTTCGGGTTTCCCCATAC 126-136 3 0.287 NED
14 SSR 863 (TC)6 55 CACAACCAGACAAGCCTTCA ACTAACGGCGGTAGCTGAGA 129-181 10 0.678 FAM
Total 79 8.379
Mean 5.6 0.598
품종간 유전적 유연관계 분석
국화 DNA profile 데이터베이스의 14개 SSR 분자표지를 이용하여 147개의 국화품종에 대한 유전적 유연관계를 알아 보고자 dendrogram을 작성하였다(Fig. 2). 공시품종의 유전 적 유사도 지수는 0.44 ~ 1.00의 범위에 분포하였고 ‘Borami (보라미)’, ‘Boramiorange(보라미오렌지)’ 와 ‘Pingpong(핑 퐁)’, ‘Whiteruuner(화이트런너)’ 품종을 제외한 143 품종이 본 연구에서 선발된 14개 SSR 분자표지의 유전자형에 따라 식별됨을 확인하였다. ‘Borami(보라미)’ 와 ‘Boramiorange (보라미오렌지)’ 품종은 유전적 유사도 지수가 100%를 나타 냈으며 이들 품종의 육성내역을 조사한 결과 ‘Boramiorange (보라미오렌지)’ 품종은 ‘Borami(보라미)’ 품종에 감마선 처 리를 하여 육성한 품종으로 형태적 특성을 조사한 결과 화색 의 형질에서만 뚜렷한 구별성을 보이는 것으로 보고된 바 있 다(Choi et al. 2014). 또한 유전적 유사도 지수가 100%인 ‘Whiterunner(화이트런너)’ 와 ‘Pingpong(핑퐁)’ 품종의 육성 내역을 알아본 결과 ‘Whiterunner(화이트런너)’ 품종은 ‘Pingpong(핑퐁)’ 품종을 재배 중 꽃이 크고 백색인 우수 변 이종을 발견하여 육성된 품종으로 확인되었다. 국화의 경우 신품종을 육성 시 돌연변이 육종방법 중 하나인 방사선 육종 법을 유용하게 이용하고 있는데 국화는 다른 종에 비해 화색 의 변이 폭이 넓고, 화형, 초장 등의 변이를 관찰할 수 있어 감마선이나 이온빔 등을 이용하여 다양한 형태를 보이는 신 품종 육성이 활발히 진행된다고 보고된 바 있다(Park & Song 2005). Shirao et al. (2013)는 국화의 품종 육성 시 변 이에 의해 육성된 품종의 경우 SSR 분자표지로 분석하였을 때 대조품종과 같은 밴드 패턴이 나타날 수 있음을 보고하였 고 Wolff & Peters-Van Rijin (1993)는 RAPD 마커로 국화 품종을 분석시 돌연변이로 육성된 품종은 분자표지로 구분되 지 않았는데 그 원인으로 형태적 변이가 나타난 부분이 국화 유전체에서 극히 작은 영역이기 때문인 것으로 보고한 바 있 다. 향후 분자표지로 구분되지 않은 품종들에 대해서는 NGS 분석을 통해 품종간 염기서열의 변이를 발견함으로써 이를 구분할 수 있는 마커를 추가 개발해야 할 것으로 사료된다.
Fig. 2.
Cluster dendrogram of the 147 chrysanthemum varieties using 14 simple sequence repeat markers. The numbers is at right side (1-147) refer to the list of varieties in Table 1. The scale at the bottom is jaccard’s coefficient of similarity.
KJBS-47-366_F2.gif
국화 147품종에 대한 dendrogram을 작성한 결과 본 연구 에서 분석된 국화 품종들은 국화 계통인 스탠다드, 스프레이, 화단, 포트 국화 별로 그룹화 되지 않았다. 카네이션의 경우에 도 스탠다드와 스프레이 계통 총 41품종에 대하여 13개 SSR 마커를 이용하여 유연관계를 분석한 결과 스탠다드와 스프레 이가 계통에 따라 분류하지 않음을 보고한 바 있다(Kimura et al. 2009). 이는 Zhang et al. (2014)에 의해 보고되었듯이 국 화의 경우 재배되고 있는 대부분의 품종이 다양한 종간의 교 잡으로부터 오랜 기간 육성되었기 때문에 유전적으로 매우 이형접합(heterozygous) 하므로 형태적으로 분류된 계통과 불일치 된 것으로 추정된다. 그러나 품종간 유사도 지수가 0.93이상으로 높게 나타난 품종들은 동일한 계통에 속하는 품종들이며, ‘Gayagold(가야골드)’ 와 ‘Gayaorange(가야오렌 지)’ 두 품종은 스프레이 국화로서 형태적 특성이 매우 유사 한 품종으로 확인되었는데 두 품종은 ‘SSR 51’ 분자표지로 구분이 되었다. 따라서 향후 분자표지의 개수를 추가하여 분 석하거나 형태적 특성과의 상관관계를 알아본다면 계통별로 그룹화되지 않은 점을 해석할 수 있을 것으로 사료된다.
UPOV의 분자표지에 관한 실무기술위원회인 분자생물학 및 생화학 실무작업반회의(BMT, Working Group on Biochemical and Molecular Techniques, and DNA-Profiling in Particular) 회의에서는 최근 분자표지 이용에 관한 technical guidelines protocol(TGP) 15 문서를 제정함으로써 품종보호 재배심사에 분자표지의 활용 의지를 보다 더 적극적 으로 표명하고 있다(UPOV 2013). BMT 회의에 보고된 국화 의 품종식별 연구결과로서 영국에서는 inter-simple sequence repeat(ISSR)와 inverse sequence-tagged repeat(ISTR)의 분 자표지를 이용하여 유사한 형태적 특성을 보이는 국화 39품종 의 식별에 활용한 사례를 보고한 바 있다(UPOV 2000). 품종 보호 출원품종수가 상대적으로 많은 화훼작물의 경우 기존품 종의 관리와 유사품종을 선정하는데 많은 품종에 대한 정보를 조사해야 하므로 분자표지를 대조품종 선정 시에 활용한다면 심사의 정밀도를 제고할 수 있을 것으로 생각된다. 본 연구로 부터 얻어진 국화 품종별 DNA profile 데이터베이스를 기반 으로 향후 실험실간 분자표지의 교차분석을 통하여 표준화된 데이터베이스를 구축한다면 품종보호권 침해 및 종자분쟁 발 생시 분자표지의 실용화가 가능할 것으로 기대된다.
본 연구는 국내에서 유통되고 있는 국화 147품종을 수집하 여 국화 품종의 식별을 위한 SSR 분자표지의 선발 및 이를 이용한 품종별 DNA profile 데이터베이스를 구축하기 위해 수행하였다. 품종식별에 적합한 분자표지를 선정하기 위해 20 개 품종을 대상으로 총 587개의 SSR 분자표지를 검정하여 다형성을 나타내는 27개의 분자표지를 선발하였다. 27개의 분자표지 중 다형성, 재현성, 반복성, 대립유전자 패턴 등을 종합적으로 고려하여 14개의 SSR 분자표지를 데이터베이스 구축에 활용할 마커로 최종 선발하였고 이를 이용하여 국화 147품종에 대한 SSR 분자표지 데이터베이스를 구축하였다. 국화 147 품종을 14개의 SSR 분자표지로 분석한 결과 대립 유전자 수는 79개, 마커별 대립유전자의 분포는 3 ~ 10개로 분포하였고, 분자표지 당 평균 대립 유전자의 수는 5.6개로 나타났다. PIC 값은 0.287 ~ 0.785 범위에 분포하였으며, 평 균값은 0.598로 분석되었다. 국화 147품종에 대한 덴드로그 램을 작성하였을 때 공시품종의 유전적 거리는 0.44 ~ 1.00 범위로 나타났으며, 147품종 중 143품종은 14개 SSR 분자표 지에 의해 식별이 되었으나, 돌연변이 육종법 또는 자연변이 를 통해 육성된 2품종은 원품종과 구분이 되지 않았다. 본 연 구에 의해 구축된 국화 품종별 SSR 분자표지 데이터베이스 는 국화 출원품종의 대조품종 선정과 품종보호권 침해 및 종 자분쟁 발생시 유용하게 활용될 수 있을 것으로 사료된다.
본 결과물은 농림축산식품부의 재원으로 농림수산식품기 술기획평가원의 생명산업기술개발사업 “작물별 형태적 특성 및 DNA 검정에 의한 품종특성의 국가표준 DB 구축”(과제번 호: 313043-3)의 지원을 받아 연구되었음.
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Construction of SSR Marker Database of Chrysanthemum Varieties Collected in Korea
Korean. J. Breed. Sci.. 2015;47(4):366-375.   Published online December 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.4.366
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Construction of SSR Marker Database of Chrysanthemum Varieties Collected in Korea
Korean. J. Breed. Sci.. 2015;47(4):366-375.   Published online December 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.4.366
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Construction of SSR Marker Database of Chrysanthemum Varieties Collected in Korea
Image Image
Fig. 1. Polymorphism of 3 simple sequence repeat markers, SSR-48, SSR-127, SSR-330. The PCR products were separated using a QIAxcel Advanced System (QIAGEN). Lane 1: ‘Borami’, 2: ‘Mujigae’, 3: ‘Hwiparam’, 4: ‘Yesmorning’, 5: ‘ Hibaeksan’, 6: ‘Magic’, 7: ‘Yellowcap’, 8: ‘Sumi’, 9: ‘Ilweol’, 10: ‘Cherryblossom’, 11: ‘Forestaroma’, 12: ‘Secretpink’, 13: ‘Baekma’, 14: ‘OrangeND’, 15: ‘PeachND’, 16: ‘BrightND’, 17: ‘PowerND’, 18: ‘Dreamround’, 19: ‘Firepink’, 20: ‘Shinma’. The A, B, C indicate specific allele type.
Fig. 2. Cluster dendrogram of the 147 chrysanthemum varieties using 14 simple sequence repeat markers. The numbers is at right side (1-147) refer to the list of varieties in Table 1. The scale at the bottom is jaccard’s coefficient of similarity.

Fig. 1.

Fig. 2.

Construction of SSR Marker Database of Chrysanthemum Varieties Collected in Korea

Chrysanthemum varieties used for genetic relationship analysis using simple sequence repeat markers.

No. Variety name Type classification No. Variety name Type classification
1 Borami Spray 45 Delidantepurple Spray
2 Mujigae Spray 46 Delibiscari Spray
3 Hwiparam Spray 47 Delitents Spray
4 Yesmorning Spray 48 Deliaragon Spray
5 Magic Spray 49 Deliquebec Spray
6 Yellowcap Spray 50 Delimarimo Spray
7 Ilweol Spray 51 Delithm Spray
8 Cherryblossom Spray 52 Lovemine Spray
9 Forestaroma Spray 53 Yeosim Spray
10 Secretpink Spray 54 Magicstar Spray
11 OrangeND Spray 55 Delmonte Spray
12 PeachND Spray 56 Moulinrouge Spray
13 BrightND Spray 57 SunnyND Spray
14 PowerND Spray 58 BounceND Spray
15 Dreamround Spray 59 Athos(Sp.) Spray
16 Firepink Spray 60 HappyND Spray
17 GraceND Spray 61 Boramiorange Spray
18 Kingfisher Spray 62 Azma Spray
19 Yesruby Spray 63 Sasineyellow Spray
20 Vogue Spray 64 Nostar Spray
21 CutieND Spray 65 Sasinecream Spray
22 Merrybell Spray 66 Caesarcream Spray
23 Dreamhappy Spray 67 Seireine Spray
24 Dreamriver Spray 68 Seiribbu Spray
25 Deliacream Spray 69 Seimyra Spray
26 Purplecap Spray 70 Seited Spray
27 Lollipop Spray 71 Seiastron Spray
28 Pinkmam Spray 72 Seielza Spray
29 Argus Spray 73 Sweetbongbong Spray
30 Purplecone Spray 74 Leopard Spray
31 Biarittz P Spray 75 Sweetsnow Spray
32 Goldrich Spray 76 Ford Spray
33 Monalisa Spray 77 Sweetsun Spray
34 Goldenboy Spray 78 Castelli Spray
35 Bacardifill Spray 79 Sweetcandy Spray
36 Pinkyboy Spray 80 Artirisingsun Spray
37 Gloria Spray 81 Pinky Spray
38 Vatican Spray 82 Artipurplelady Spray
39 Yeonja Spray 83 Artidarkchocolate Spray
40 Naos Spray 84 Yesholic Spray
41 Sasine Spray 85 Jazz Spray
42 Neseteo Spray 86 Yellowdream Spray
43 Seifrillgreen Spray 87 Spybelly Spray
44 Delibarca Spray 88 Yellowmarble Spray
89 Seiagness Spray 119 Zemblalime Standard
90 Yellowkid Spray 120 Armadolis Pot mum
91 Champy Spray 121 Aamflu Pot mum
92 Orangedream Spray 122 Myruby Pot mum
93 Zarita Spray 123 Vino Pot mum
94 Orangekid Spray 124 Mygirl Pot mum
95 Hirario Spray 125 Pinkeye Pot mum
96 Zembla Spray 126 Mysoul Pot mum
97 Cutiepink Spray 127 Gayagold Pot mum
98 Cheeks Spray 128 Gayaorange Pot mum
99 Fireyellow Spray 129 Gayaglory Pot mum
100 Karisyellow Spray 130 Gayayellow Pot mum
101 Pinkycharis Spray 131 Gayaneon Pot mum
102 Yellowpangpang Spray 132 Gayaruby Pot mum
103 Pinkpangpang Spray 133 Purpleegg Pot mum
104 Rubyna Spray 134 Juliet Pot mum
105 Hongpa Spray 135 Peacecutie Pot mum
106 Hibaeksan Standard 136 Gayapink Pot mum
107 Sumi Standard 137 Mygold Pot mum
108 Baekma Standard 138 Mysunny Pot mum
109 Shinma Standard 139 Myred Pot mum
110 Geumhwa Standard 140 Gayasense Pot mum
111 Boryeonghwangkuk Standard 141 Mywing Pot mum
112 GreenND Standard 142 Mycream Pot mum
113 Athos(St.) Standard 143 Candyball Garden mum
114 Deliakari Standard 144 Dasomball Garden mum
115 Whiterunner Standard 145 Maruball Garden mum
116 Pingpong Standard 146 Madangball Garden mum
117 Seolmi Standard 147 Geumbangul Garden mum
118 Ecogreen Standard

Simple sequence repeat markers screened for identifying polymorphism of chrysanthemum.

Number of screened markers Type of SSR markers Number of polymorphic markers Polymorphism (%) of amplified SSR markers SSR marker source
12 Genomic SSR of chrysanthemum 2 2/12 Khaing et al. (2013)
575 EST-SSR of chrysanthemum 25 25/575 Wang et al. (2013)
587 27 27/587 (4.6%)

Characteristics of the 14 simple sequence repeat markers selected for genetic characterization of chrysanthemum varieties.

No. Marker name Repeat motif Annealing temp. (°C) Forward sequence Reverse sequence PCR product size (bp) No. of alleles PIC value Primer labeling
1 SSR 48 (CCA)5 55 TGAGATCATTCCCAACCTCC CTAGCGTCCAAAGAATTGGC 206-214 4 0.655 VIC
2 SSR 51 (CCAA)4 55 CCCCCTCTTCTTCTTCAACC CAATAGAAAGCGCGTGACAA 197-252 8 0.785 NED
3 SSR 52 (ATA)5 55 AGTGACCCGAGCCAGATAGA CCGACAAATCATTTCCGTCT 233-247 6 0.573 FAM
4 SSR 127 (GGT)5 55 TAACAAGGGGTTTCAGCGTC TCAGGAAGAACAACCCAACC 262-273 5 0.335 FAM
5 SSR 200 (GGT)5 55 CCCAGAGAAGCGTGAGATTT TCCCCTGCTACTACCACCAC 129-154 4 0.637 VIC
6 SSR 222 (ATC)5 55 AGCTAAAACAAACAAGCGGC GCGTTAACTGTGTCGGTTGA 153-185 8 0.756 NED
7 SSR 330 (CAA)5 55 CTGTTGAGCAGTTCAGGCAC GTGTGATCGAGGCGATTTTT 204-219 6 0.597 FAM
8 SSR 332 (TGG)5 55 ACACCGAATAGGACGGACAG TTTTCTGAAGTCCCGACCAC 227-240 5 0.556 PET
9 SSR 380 (TGA)5 55 ACCAAAGGCAGCTCACAGTT CCTCCCTCACTCATCTCTGC 190-196 4 0.658 VIC
10 SSR 649 (ACC)7 55 TCTTCCTCACACGCAAACAC AGCTGCCACTCGCTATCACT 247-259 6 0.762 FAM
11 SSR 706 (AG)8 55 CGATCACCATTCTTTTCCCA CCGATAAGTTCGTCCTTGGT 208-240 7 0.613 PET
12 SSR 728 (ATG)5 55 TGGTTATGGGTGACCCTGAT AAGAAAGTGCAGGCCAAGAA 145-151 3 0.488 VIC
13 SSR 792 (TGA)6 55 AGGAAGAAGATCGACACCCA AAGTTCGGGTTTCCCCATAC 126-136 3 0.287 NED
14 SSR 863 (TC)6 55 CACAACCAGACAAGCCTTCA ACTAACGGCGGTAGCTGAGA 129-181 10 0.678 FAM
Total 79 8.379
Mean 5.6 0.598
Table 1. Chrysanthemum varieties used for genetic relationship analysis using simple sequence repeat markers.
Table 2. Simple sequence repeat markers screened for identifying polymorphism of chrysanthemum.
Table 3. Characteristics of the 14 simple sequence repeat markers selected for genetic characterization of chrysanthemum varieties.

Table 1.

Table 2.

Table 3.

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