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-allele Specific PCR 분석에 의한 사과와 꽃사과 품종의 자가불화합성 유전자형 동정

Identification of Self-Incompatibility Genotypes in Apple and Crabapple Cultivars by S-allele Specific PCR Analysis

Kang Hee Cho1,†,*, Jeong-Hee Kim2,†, Jung Woo Lee2, Soon-Il Kwon2, Jong Taek Park2, Il Sheob Shin1, Se Hee Kim1, Dae-Hyun Kim1, In Myeong Choi1
Korean Journal of Breeding Science 2014;46(4):364-371.
Published online: October 31, 2014

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*Corresponding Author : (khc7027@korea.kr, Tel: +82-31- 240-3676, Fax: +82-31-240-3709)

The first two authors contributed equally to this work.

• Received: September 19, 2014   • Revised: October 16, 2014   • Accepted: October 17, 2014

© Tshe Korean Society of Breeding Science All rights reserved

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  • Apple (Malus × domestica Borkh.) has gametophytic self-incompatibility (S) controlled by the multi-allelic S-locus. In the present study, S-genotypes of 24 apple cultivars including newly released Korean cultivars and seven crabapple cultivars were identified using S-allele specific polymerase chain reaction analysis. Twelve different S-alleles (S1, S2, S3, S5, S7, S9, S10, S16, S21, S23, S26, and S29) from 31 apple and crabapple cultivars were identified using 23 S-allele specific primers. Among them, S1 (41.7%), S3 (58.3%), S7 (29.2%), and S9 (54.2%) S-alleles were found to be common in 24 apple cultivars. The newly released Korean cultivars ‘Arisoo’ and ‘Hwangok’ were genetyped as S3S7 and S3S9, respectively. S-genotypes information obtained from the present study will be useful to select proper pollinzers for stable production of apple fruit and to design cross of breeding programs.
사과는 배우체형 자가불화합성(gametophytic self-incompatibility) 작물로, 자가불화합성은 단일유전자 좌인 S-locus의 복대립유전자에 의해 조절된다(Kobel et al. 1939). S-locus 의 주두측 산물은 ribonuclease의 활성을 갖는 당단백질 (glycoprotein)인 S-RNase로 화분측의 자가불화합성 유전자 인 F-box와의 상호작용에 의해 자가수분을 억제하는 것으로 알려져 있다(Hegedüs 2006). 따라서 안정적인 결실을 위해서 는 다른 품종의 꽃가루로 인공수분을 시키거나, 수분수 전용 꽃사과 품종 혹은 자가불화합성 유전자형(S-genotype)이 다 른 재배품종을 혼식해야 한다. 기존의 과수원은 대부분 재배 품종을 수분수로 이용하고 있으나 과실 수확시기, 병해충 발 생 정도 등이 서로 달라 관리상 어려운 점이 많다. 따라서 사 과 품종의 S-genotype 정보는 새로운 품종의 육종과 과실 생 산을 위한 수분수 선발에 매우 중요하다(Kim et al. 2008).
사과의 S-genotype을 동정하기 위한 다양한 연구들이 진행 되어 왔다. Kobel et al. (1939)은 26품종을 이용하여 다양한 교배 후 화분관 신장 및 결실율을 조사하여 11개의 S-allele (S1-S11)을 동정하였다. 그러나 교배를 통해 자가불화합성을 결정하는 방법은 시간과 노력이 많이 소요되고, 화분관 신장 이 환경과 생리적 조건에 영향을 받아 결과가 명확하지 않은 경우가 있다(Van Nerum et al. 2001). Broothaerts et al. (1995)과 Sassa et al. (1996)은 S-RNase에 암호화하는 cDNA 를 분리하였고, 장미과 S-RNase의 염기서열에는 5개의 보존 영역(conserved region: C1-C5)과 C2와 C3사이에 하나의 변 이영역(hypervariable region)이 존재하는 것으로 밝혀졌으며, 변이영역 내에 포함된 인트론은 다형성을 나타낸다(Wu et al. 2013). 이러한 S-RNase의 염기서열 정보를 기초로 하여 S-allele을 동정할 수 있는 분자생물학적 기술이 개발되었다. S-genotype 결정하는 방법에는 S-allele specific primer를 이 용한 polymerase chain reaction (PCR) 분석과 제한효소를 이용하는 restriction fragment length polymorphism 분석이 있다(Broothaerts 2003, Janssens et al. 1995, Kim et al. 2008, Kitahara & Matsumoto 2002, Long et al. 2010). Van Nerum et al. (2001)은 Boskovic & Tobutt (1999)이 단백질 분석에 의해 동정된 사과 56품종의 S-genotype을 S-allele specific PCR과 염기서열 분석을 통하여 14품종의 S-genotype 을 재정립하였고, Broothaerts (2003)은 사과의 자가불화합성 유전자들을 새롭게 15 S-allele (S1, S2, S3, S4, S5, S7, S9, S10, S16, S19, S20, S22, S23, S24, S26)로 정립하였다. Kim et al. (2008)은 ‘York Imperial’과 ‘Burgundy’ 품종으로부터 각각 S31과 S32의 S-allele을 동정하였고, Long et al. (2010)은 S44, S45, S46의 새로운 S-allele을 보고하였다.
국내에서 교배를 통한 사과 품종 육종은 1988년 국립원예 특작과학원에서 육성한 ‘홍로’ 품종을 시작으로 현재까지 꽃 사과 3종을 포함하여 27품종이 육성되었다. ‘후지’ 등 기존 품종들의 S-genotype은 밝혀져 있지만 ‘아리수’, ‘썸머킹’ 등 국내에서 육성한 신품종은 S-genotype에 대한 정보가 잘 알 려져 있지 않다. 새로 육성된 사과 품종들의 정확한 S-allele 의 구명은 기존 주요 품종과 신품종과의 수분 가능 여부 등을 판단하는데 매우 중요하다.
따라서 본 연구는 S-allele specific PCR분석을 통해 신품 종을 포함한 사과 주요 재배품종과 꽃사과 품종의 S-genotype 을 결정하여 육종프로그램에서 교배조합 작성, 품종재배 시 최적 수분수 선정 등의 기초자료로 활용하고자 하였다.
시험재료 및 Genomic DNA 추출
국립원예특작과학원 사과시험장에 재식되어 보존중인 사 과 국내육성 품종 17종, 도입품종 7종의 재배품종과 최근 개 발된 ‘골든벨’과 ‘로즈벨’ 품종이 포함된 7종의 꽃사과 품종 을 실험재료로 사용하였다(Table 1). 사과나무의 어린 잎을 채취하고 액체질소로 곱게 마쇄한 후 DNeasy plantmini kit (Qiagen, Valencia, CA, USA)를 사용하여 genomic DNA를 추출하였다. 추출한 DNA의 순도는 0.8% agarose gel에 전 기영동하여 확인하였고 DNA 양은 분광광도계(NanoDrop 2000, Thermo Fisher Scientific, MA, USA)로 정량분석하 였다. PCR분석에는 DNA를 5 ng·μL-1의 농도로 희석하여 이 용하였다.
Table 1.
List of apple and crabapple cultivars used in this study.
Table 1.
No. Cultivar Parentage (S-genetype) Referencez Origin
Apple
1 Alps Otome (S1S9) Unknown - Japan
2 Arisoo (S3S7) Yoko (S3S9) × Senshu (S1S7) 1, 5 Korea
3 Chukwang (S3S9) Fuji (S1S9) × Mollie’s Delicious (S3S7) 1, 2, 3, 4, 5 Korea
4 Fuji (S1S9) Ralls Janet (S1S2) × Delicious (S9S19) 1, 2, 5 Japan
5 Gala (S2S5) Kidd's Orange Red (S5S9) × Golden Delicious (S2S3) 1, 2, 5 USA
6 Gamhong (S1S9) Spur Earliblaze (S1S19) × Spur Golden Delicious (S2S3) 4 Korea
7 Green Ball (S3S7) Golden Delicious (S2S3) × Fuji (S1S9) 1, 2, 5 Korea
8 Hongan (S1S3) Fuji (S1S9) × Jonathan (S7S9) 1, 2, 5 Korea
9 Honggeum (S3S7) Senshu (S1S7) × Hongro (S1S3) 1, 2, 4, 5 Korea
10 Hongro (S1S3) Spur Earliblaze (S1S19) × Spur Golden Delicious (S2S3) 4 Korea
11 Hongso (S1S3) Yoko (S3S9) × Hongro (S1S3) 3, 4, 5 Korea
12 Hwahong (S3S9) Fuji (S1S9) × Sekaiichi (S3S9) 1, 5 Korea
13 Hwangok (S3S9) Kogetsu (S3S7) × Yataka Fuji (S1S9) 1 Korea
14 Manbok (S1S9) Earliblaze (S1S19) × Fuji (S1S9) 1, 4, 5 Korea
15 Picnic (S1S5) Fuji (S1S9) × Sansa (S5S7) 1, 4, 5 Korea
16 Sansa (S5S7) Gala (S2S5) × Akane (S7S24) 1, 4, 5 Japan
17 Seohong (S3S9) Tsugaru (S3S7) × Chukwang (S3S9) 1, 2, 3, 4, 5 Korea
18 Sinano Sweet (S1S7) Fuji (S1S9) × Tsugaru (S3S7) 1, 2, 3, 5 Japan
19 Summer Dream (S7S9) Tsugaru (S3S7) × Natsumidori (S3S9) 1, 2, 3, 5 Korea
20 Summer King (S3S9) Fuji (S1S9) × Golden Delicious (S2S3) 1, 2, 5 Korea
21 Sunhong (S3S9) Hongro (S1S3) × Chukwang (S3S9) 3, 4 Korea
22 Tsugaru (S3S7) Golden Delicious (S2S3) × Unknown 1, 2, 5 Japan
23 Yeohong (S1S9) Jonathan (S7S9) × Fuji (S1S9) 1, 2, 5 Korea
24 Yoko (S3S9) Golden Deliciou s(S2S3) × Unknown 1, 2, 5 Japan
Crabapple
25 Golden Bell (S2S23) Gala (S2S5) × Mo.84 (S4S26) 1, 2, 5 Korea
26 Malus prunifolia (S7S29) Wild species China
27 Maypole (S10S16) McIntosh Wijcik (S10S25) × Malus Baskatong (S16S26) 1, 2 France
28 Purple Lemoine (S7S9) Malus × atrosanguinea × Malus niedzwetskyana - France
29 Rose Bell (S21S26) Gala (S2S5) × Mo.84 (S4S26) 1, 2, 5 Korea
30 Sentinel (S21S26) Unknown - New Zealand
31 Yantaishagou (S9S21) Unknown - -

zParentage’s S-genetypes were reported by references. 1, Broothaerts (2004); 2, Heged s (2006); 3, Heo et al. (2011); 4, Kim et al. (2006); 5, Yamane & Tao (2009)

S-allele specific primer를 이용한 PCR분석
사과 재배품종과 꽃사과 품종의 S-genotype을 동정하기 위 하여 기존에 보고된 Broothaerts (2003), Long et al. (2010) 과 Son et al. (2013)의 21종의 S-allele specific primer 를 본 실험에 사용하였다(Table 2). PCR 반응은 총 15 μL 반응액 에 genomic DNA 40 ng, 200 μM dNTP, 5 pmol S-allele specific primer, 1 × PCR buffer (Genetbio, Daejeon, Korea) 및 PCR 반응 시 비특이적 DNA 증폭을 최소화하기 위하여 0.5 unit의 Hot-start Taq DNA polymerase를 첨가하여 수행하였 다. PCR 기기는 C-1000 thermocycler (Bio-Rad Laboratories, Hercules, CA, USA)를 사용하여, 95°C에서 10분간 초기변 성 후 95°C에서 30초, 55-66°C에서 30초, 72°C에서 1분간 총 30회 반복하고 72°C에서 5분간 처리한 후 반응을 종료하였 다. PCR 산물은 1.4% agarose gel에서 전기영동하여 S-genotype 을 확인하였다. S-allele S16과 S22의 동정은 FTC5, OWB249 primer를 이용하여 PCR 수행 후 증폭산물을 TaqI 제한효소 로 65°C에서 2시간 처리하였고, 3% agarose gel에 전기영동 하여 절단된 단편 수와 크기에 따라 S-genotype을 결정하였다.
Table 2.
S-allele specific primers used for PCR amplification.
Table 2.
S-allele Primer Sequence (5’→3’) Amplicon size (bp) Annealing temp. (°C) Reference
S1 MdS1SpF TGTAAGGCACCGCCATATCATAC 734 62 Long et al. (2010)
MdS1SpR CAACCTCAACCAATTCAGTCAATGA
S2 MdS2SpF AACATGAATCGAAGTGAATTATTTA 489 55 ”
MdS2SpR TTGAGGTTTGGTTCCTTACCATG
S3 MdS3SpF GGCGAAAATTAAACCGGAGAAGAA 292 58 ”
MdS3SpR CCTCTCGTCCTATATATGGAAATCAC
S4 MdS4SpF ATTGCAAGACAAGGAATCGTCGGAA 533 63 ”
MdS4SpR AGAAATGTGCTCTGTTTTTATCG
S5 FTC10 CAAACATGGCACCTGTGGGTCTCC 346 60 Broothaerts (2003)
FTC11 TAATAATGGATATCATTGGTAGG
S7 MdS7SpF AGTAAATCAACCGTGGATGCTCAG 397 63 Long et al. (2010)
MdS7SpR TTACAATATCTACCTGTTTCCTGGG
S9 MdS9SpF CCACTTTAATCCTACTCCTTGTAGA 522 63 ”
MdS9SpR TCAATTTCCTTCTGTGTCCTGAATT
S10 FTC12 CCAAACGTACTCAATCGAAG 209 66 Broothaerts (2003)
FTC228 ATGTCGTCCCGTGTCCTGAATC
S11 MdS11SpF AAATATTGCAAGGCGCCGC 678 63 Long et al. (2010)
MdS11SpR TTTCAATATCTACCAGTCTCCGGC
S16 FTC5 TCCCACAATACAGAACGAGA 274, 243+31 (TaqI) 60 Broothaerts (2003)
OWB249 CAATCTATGAAATGTGCTCTG
S19 MdS19SpF GCCTTCAAACAAGAATGGACC 481 63 Long et al. (2010)
MdS19SpR TCAATATCCACCAATGACCTGTT
S20 MdS20SpF GTTGTGGCCTTCAGACTCG 882 64 Long et al. (2010)
MdS20SpR GGCCAACTACTTTTATTTTTCATC
S21 MdS21SpF AAGTAATTGCCCGATAAGGAACATA 584 63 Long et al. (2010)
MdS21SpR AGTTTATGAAATGTTCTCCGCTGTA
S23 FTC222 CAATCGAACCAATCATTTGGT 237 60 Broothaerts (2003)
FTC224 GGTGTCATATTGTTGGTACTAATG
S24 MdS24SpF ATGGCTCCTGTGCGTCTTCCC 421 61 Long et al. (2010)
MdS24SpR CGTCATCCGTGTATAGGGCAACT
S26 MdS26SpF TCCATCAAACGTGACTTCTCAT 423 55 ”
MdS26SpR ATCCTTCAGCATCCTGATTCG
S29 A29PL TATAAACATGGCTCCTGTGCG 217 60 Son et al. (2013)
A29PR TCGTTTGGCACTTGAGTTTTG
S31 MdS31SpF TGACCCAAAATATTGCAAGGCGC 556 63 Long et al. (2010)
MdS31SpR TTTCAATATCTACCAGTCTCCGGC
S44 MdS44SpF GCATGGTAGGACCTGACCCAAGTA 561 63 ”
MdS44SpR TCTCAACCAATTGAGTCGTCGTACC
S45 MdS45SpF CCAGAAGGTTGCAAGACACAGAAAT 641 63 ”
MdS45SpR AGTTTTGGTGCCTTATTGTTGGTAC
S46 MdS46SpF CCCAACGTGCTCGATCGAACA 212 61 ”
MdS46SpR TCAATTTCCTTCTGTGTCCAGAATC
사과 신품종의 S-genotype에 대한 정보는 신육성 품종의 재식 시 주요 품종과의 교배 친화성 여부를 사전에 판단할 수 있는 정보를 제공하기 때문에 이를 바탕으로 품종 구성을 선 택하면 사전에 결실 불량 요인을 차단할 수 있다. 이는 신육 성 품종 보급률을 높이고 안정적인 결실 및 정형과 생산에 매 우 중요하다. 본 실험에서는 새로 육성한 품종 ‘아리수’, ‘황 옥’ 등을 포함한 주요 재배품종과 꽃사과 품종의 S-genotype 을 구명하고자 하였다.
사과 재배품종 24종과 꽃사과 7종의 총 31품종의 유엽에 서 추출한 DNA와 S-allele specific primer 를 이용하여 PCR 분석을 수행하였다. 그 결과 S1, S2, S3, S5, S7, S9, S10, S16 (S22), S21, S23, S26, S29의 12개 S-allele이 증폭되었고(Fig. 1), S4, S11, S19, S20, S24 S31, S44, S45, S46 은 증폭되지 않았다. MdS1Sp primer를 이용한 PCR 분석에서는 31품종 중에서 ‘알프스오또메’, ‘후지’, ‘감홍’, ‘홍안’ 등 10종의 재배품종에 서만 734 bp크기의 S1 allele이 증폭되었고, MdS3Sp primer 를 이용한 분석에서는 ‘아리수’, ‘추광’, ‘그린볼’, ‘썸머킹’ 등 14종의 재배품종에서만 292 bp크기의 S3 allele이 증폭되었 다(Fig. 2). 본 실험에서 증폭된 S-allele의 크기는 염기서열 분석을 통하여 정확히 확인하지 않았기 때문에 기존에 논문 에서 보고된 크기를 인용하여 나타내었다. S2 allele은 ‘갈라’ 와 ‘골든벨’ 품종에서만 489 bp크기의 밴드로 동정되었고, S5 allele은 ‘피크닉’, ‘갈라’, ‘산사’ 품종에서만 346 bp 크기의 밴드로 동정되었다. S5 allele은 Long et al. (2010)이 보고한 MdS5Sp primer를 이용하여 분석하였을 때는 Heo et al. (2011)의 결과와 동일하게 증폭되지 않았다. 55~64°C의 다 양한 annealing 온도조건에서도 증폭이 되지 않았는데 이는 S10 과 S23 allele을 증폭하는 primer도 동일하였다. 이러한 결 과는 S-allele을 가진 품종의 염기서열에 single nucleotide polymorphism이 존재하는데 기인하는 것으로 추정하였다 (Heo et al. 2011). 따라서 이들 S-allele의 다형성을 회복하기 위하여 많은 노력이 필요하므로 본 실험에서는 Broothaerts (2003)이 보고한 primer를 이용하여 S5, S10, S23 allele을 동 정하였다(Table 2).
Fig. 1.
Genomic PCR amplification products using specific primers for each of the S-alleles in apple. M, 100 bp plus DNA ladder (Bioneer); S1, ‘Hongan’; S2, ‘Gala’; S3, ‘Arisoo’; S5, ‘Sansa’; S7, ‘Honggeum’; S9, ‘Summer King’; S10, ‘Maypole’; S16 (S20), ‘Maypole’; S21, ‘Rose Bell’; S23, ‘Golden Bell’; S26, ‘Sentinel’; S29, Malus prunifolia.
KJBS-46-364_F1.gif
Fig. 2.
Results of PCR amplification with MdS3SpF and MdS3SpR primers in 31 apple cultivars. Cultivars correspond to the 31 genotypes listed in Table 1. M, 100 bp plus DNA ladder.
KJBS-46-364_F2.gif
S-allele specific primer를 이용한 사과 31품종의 S-genotype 분석 결과는 Table 3에 나타내었고, ‘썸머킹’, ‘황옥’ 등 신육 성 품종을 포함한 24종 재배품종의 S-genotype 분석 결과는 Fig. 3과 같다. S1S3 의 S-allele 을 가진 품종은 ‘홍안’, ‘홍 로’, ‘홍소’ 로, ‘홍로’ 품종의 S-genotype은 모본의 ‘스퍼어리 브레이즈’(S1S19)와 부본의 ‘스퍼 골든 데리셔스’(S2S3)로부터 유래되었음을 확인할 수 있었다. 그러나 ‘홍안’ 품종은 ‘후지’ (S1S9)와 ‘홍옥’(S7S9)을 교배하여 육성된 품종이나 S-genotype 은 S1S3 로 나타났으며, ‘골드 데리셔스’(S2S3)와 ‘후지’(S1S9) 를 교배하여 육성된 품종인 ‘그린볼’은 S3S7 로 확인되었다. ‘감홍’(S1S9) 품종도 양친인 ‘스퍼어리 브레이즈(S1S19)와 ‘스 퍼 골든 데리셔스’(S2S3)품종에는 S9 allele이 존재하지 않았 다. 이들 품종들은 교배 시 다른 품종의 화분이 이용되었거나 방임수분 등에 기인한 것으로 추정되며 추후 양친에 대한 재 검토가 필요하다(Heo et al. 2011). 중과종으로 육성된 ‘피크 닉’의 S-genotype은 S1S5로 확인되어 ‘후지’와 ‘홍로’를 포함 한 대부분의 재배품종과 교배친화성이 높아 수분수로 이용이 가능하다. ‘시나노 스위트’와 ‘산사’ 품종의 S-genotype은 각 각 S1S7과 S5S7로 확인되었다. S3S7의 S-genotype은 ‘그린볼’ 을 포함하여 국내육성 품종인 ‘아리수’, ‘홍금’과 도입품종인 ‘쓰가루’에서 확인되었다. ‘아리수’는 ‘양광’(S3S9)과 ‘천추’ (S1S7)를 교배하여 2010년에 육성된 품종으로 착색기에 기온 이 높은 지역에서도 착색이 잘 되는 장점이 있다. S3S9 의 S-genotype은 신육성 품종인 ‘황옥’과 ‘썸머킹’ 등 6종의 국 내육성 품종과 도입품종인 ‘양광’에서 확인되었고, ‘쓰가루’ (S3S7)와 ‘하록’(S3S9) 품종을 교배하여 육성된 ‘썸머드림’ 품 종은 S7S9 의 S-genotype을 나타내었다. 이와 같이 국내 육성 품종에서는 주로 S1(41.7%), S3(58.3%), S7(29.2%), S9(54.2%) 의 S-genotype이 확인되었는데 이는 국내 사과 육종프로그램 에서 양친으로 대부분 ‘후지’(S1S9), ‘쓰가루’(S3S7), ‘홍옥’ (S7S9), ‘스퍼 골든 데리셔스’(S2S3)와 같은 소수의 외국품종 이 사용되었기 때문으로 추정된다. 따라서 국내에서 육성된 사과 품종의 유전적 변이의 폭은 좁은 것으로 판단되므로 새 로운 품종을 육성하기 위해서는 다양한 유전자원을 육종프로 그램에 도입하여 변이의 폭을 확대할 필요가 있다.
Fig. 3.
Genotyping of S-allele by PCR with specific primers in 24 apple cultivars. The PCR product sizes were reported by Broothaerts (2003) and Long et al. (2010). Cultivars correspond to the 24 genotypes listed in Table 1. M, 100 bp plus DNA ladder.
KJBS-46-364_F3.gif
Table 3.
S-allele genotypes of apple and crabapple cultivars by S-allele-specific PCR.
Table 3.
S-allele genotype Cultivar
S1S3 Hongan, Hongro, Hongso
S1S5 Picnic
S1S7 Sinano Sweet
S1S9 Alps Otome, Fuji, Gamhong, Manbok, Yeohong
S2S5 Gala
S2S23 Golden Bell
S3S7 Arisoo, Green Ball, Honggeum, Tsugaru
S3S9 Chukwang, Hwahong, Hwangok, Seohong, Summer King, Sunhong, Yoko
S5S7 Sansa
S7S9 Summer Dream, Purple Lemoine
S7S29 Malus prunifolia
S9S21 Yantaishagou
S10S16 Maypole
S21S26 Rose Bell, Sentinel
꽃사과는 해에 따라 개화기 차이가 있으므로 수분수로 사 용 시 3~4품종을 혼식하는 것이 안전하다. 재배품종의 수분 수로 활용하기 위하여 7종의 꽃사과 품종의 S-genotype을 분 석 결과는 Fig. 4와 같다. ‘Purple Lemoine’ 품종은 ‘썸머드 림’과 동일한 S7S9 로, Malus prumifolia는 S7S29의 S-allele 이 확인되었다. ‘Purple Lemoine’와 Malus prumifolia는 기 존의 보고에서는 S-genotype을 동정할 수 없었으나(Son et al. 2013), 본 실험을 통하여 이들의 S-genotype이 밝혀졌다. 또한 기존 보고에서 S26의 S-allele 한가지 유전자형만 동정되 었던 ‘Sentinel’ 품종은 S21S26으로, S9 S-allele만 동정되었던 ‘Yantaishagou’ 품종은 S9S21로 확인되었다. S16과 S22 S-allele 의 염기서열은 99%가 동일하여 각각의 allele을 증폭할 수 있 는 특이적인 primer를 제작하기 어렵지만, 이들의 allele은 제 한효소 TaqI이 인지하는 염기서열 부위에 의해 구분할 수 있 다(Broothaerts 2003, Long et al. 2010). S16과 S22 S-allele 을 증폭할 수 있는 특이적인 FTC5와 WB249 primer를 이용 하여 PCR분석 후 증폭된 274 bp크기의 PCR산물을 TaqI으 로 처리하여 전기영동으로 확인하면 S16 allele은 243 bp와 31 bp크기의 밴드를, S22 allele은 199 bp, 44 bp, 31 bp 크기 의 밴드를 갖는다(Broothaerts 2003). 본 실험에서는 FTC5와 WB249 primer를 이용한 PCR분석에서 31품종 중 ‘Maypole’ 품종만이 증폭되었고 PCR산물의 TaqI 제한효소 처리를 통하 여 243 bp와 31 bp크기의 밴드를 확인하였다. 따라서 ‘McIntosh’ (S10S25)의 무측지성 변이품종인 ‘McIntosh Wijcik’과 꽃사과 품종인 ‘Baskatong’의 교배를 통해 육성된 ‘Maypole’ 품종의 S-genotype은 S10S16으로 확인되어 Van Nerum et al. (2001) 의 결과와 일치하였다. ‘Baskatong’ 품종의 S-genotype은 S16S26으로 알려져 있다(Broothaerts 2003, Verdoodt et al. 1998). ‘골든벨’과 ‘로즈벨’은 국립원예특작과학원 사과시험 장에서 육성한 꽃사과 품종으로 본 실험 결과 S-genotype은 각각 S2S23과 S21S26으로 확인되었다(Table 3, Fig. 4). 이 두 품종은 ‘갈라’ 품종과 대목인 Mo.84를 교배하여 육성된 품종 으로 양친이 동일하다. ‘갈라’ 품종의 S-genotype은 S2S5이나 부본으로 이용된 Mo.84의 S-genotype은 알려진 바 없다. Mo.84의 S-genotype 을 확인하기 위하여 21종의 primer를 분석한 결과 S4S26인 것으로 확인되어(자료 미제시), 추후 ‘골 든벨’과 ‘로즈벨’ 품종의 양친에 대한 검토가 필요할 것으로 판단된다. 본 실험에서는 S-allele specific PCR 분석을 통하 여 사과 31 품종 모두에서 자가불화합성 두 개의 대립유전자 에 해당되는 프라이머 조합에서만 증폭이 되었는데 이는 분석 방법의 타당성을 간접적으로 나타낸 것이다. S-allele specific primer를 이용한 분석은 S-genotype의 동정뿐 만 아니라 계 통분석 및 품종간의 유전적 변이 등을 구명하는 데에도 효율 적으로 활용될 수 있을 것으로 판단된다.
Fig. 4.
Genotyping of S-allele by PCR with specific primers in 7 crabapple cultivars. The PCR product sizes were reported by Broothaerts (2003) and Long et al. (2010). M, 100 bp plus DNA ladder. 1, Golden Bell (S2S23); 2, Malus prunifolia (S7S29); 3, Maypole (S10S16); 4, Purple Lemoine (S7S9); 5, Rose Bell (S21S26); 6, Sentinel (S21S26); 7, Yantaishagou (S9S21).
KJBS-46-364_F4.gif
현재까지 보고된 S-allele specific primer를 이용하여 분석 한 결과 다수의 꽃사과 품종의 S-genotype을 동정하기는 어 려웠다. 꽃사과 ‘Manchurian’과 ‘Spectabilis’ 품종은 각각 S24와 S45의 하나의 S-allele만 동정되었다. 따라서 추가적인 S-allele specific primer의 검토와 새로운 S-genotype 의 동 정이 가능하도록 지속적인 연구가 필요할 것으로 판단된다.
사과는 배우체형 자가불화합성을 나타내는데 이는 S-locus 의 복대립유전자에 의해 조절된다. 본 연구는 S-allele specific PCR분석을 통해 신품종을 포함한 24종의 사과 주요 재배품 종과 7종의 꽃사과 품종의 자가불화합성 유전자형(S-genotype) 을 결정하고자 수행하였다. 31종의 재배품종과 꽃사과 품종 을 23종의 S-allele specific primer을 이용하여 분석한 결과 12개의 S-allele (S1, S2, S3, S5, S7, S9, S10, S16, S21, S23, S26, S29)이 동정되었다. 그 중에서 24종의 재배품종에는 S1(41.7%), S3(58.3%), S7(29.2%), S9(54.2%)의 S-allele이 흔히 존재하 는 것으로 확인되었다. 국내육성 신품종인 ‘아리수’와 ‘황옥’ 의 S-genotype은 각각 S3S7과 S3S9으로 동정되었다. 본 실험 에서 얻은 S-genotype 정보는 안정적인 사과 과실생산에 적 합한 수분수 선발과 육종프로그램에서 교배조합 작성에 유용 하게 활용될 것이다.
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Identification of Self-Incompatibility Genotypes in Apple and Crabapple Cultivars by S-allele Specific PCR Analysis
Korean. J. Breed. Sci.. 2014;46(4):364-371.   Published online December 31, 2014
DOI: https://doi.org/10.9787/KJBS.2014.46.4.364
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Identification of Self-Incompatibility Genotypes in Apple and Crabapple Cultivars by S-allele Specific PCR Analysis
Korean. J. Breed. Sci.. 2014;46(4):364-371.   Published online December 31, 2014
DOI: https://doi.org/10.9787/KJBS.2014.46.4.364
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Identification of Self-Incompatibility Genotypes in Apple and Crabapple Cultivars by S-allele Specific PCR Analysis
Image Image Image Image
Fig. 1. Genomic PCR amplification products using specific primers for each of the S-alleles in apple. M, 100 bp plus DNA ladder (Bioneer); S1, ‘Hongan’; S2, ‘Gala’; S3, ‘Arisoo’; S5, ‘Sansa’; S7, ‘Honggeum’; S9, ‘Summer King’; S10, ‘Maypole’; S16 (S20), ‘Maypole’; S21, ‘Rose Bell’; S23, ‘Golden Bell’; S26, ‘Sentinel’; S29, Malus prunifolia.
Fig. 2. Results of PCR amplification with MdS3SpF and MdS3SpR primers in 31 apple cultivars. Cultivars correspond to the 31 genotypes listed in Table 1. M, 100 bp plus DNA ladder.
Fig. 3. Genotyping of S-allele by PCR with specific primers in 24 apple cultivars. The PCR product sizes were reported by Broothaerts (2003) and Long et al. (2010). Cultivars correspond to the 24 genotypes listed in Table 1. M, 100 bp plus DNA ladder.
Fig. 4. Genotyping of S-allele by PCR with specific primers in 7 crabapple cultivars. The PCR product sizes were reported by Broothaerts (2003) and Long et al. (2010). M, 100 bp plus DNA ladder. 1, Golden Bell (S2S23); 2, Malus prunifolia (S7S29); 3, Maypole (S10S16); 4, Purple Lemoine (S7S9); 5, Rose Bell (S21S26); 6, Sentinel (S21S26); 7, Yantaishagou (S9S21).

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Identification of Self-Incompatibility Genotypes in Apple and Crabapple Cultivars by S-allele Specific PCR Analysis

List of apple and crabapple cultivars used in this study.

No. Cultivar Parentage (S-genetype) Referencez Origin
Apple
1 Alps Otome (S1S9) Unknown - Japan
2 Arisoo (S3S7) Yoko (S3S9) × Senshu (S1S7) 1, 5 Korea
3 Chukwang (S3S9) Fuji (S1S9) × Mollie’s Delicious (S3S7) 1, 2, 3, 4, 5 Korea
4 Fuji (S1S9) Ralls Janet (S1S2) × Delicious (S9S19) 1, 2, 5 Japan
5 Gala (S2S5) Kidd's Orange Red (S5S9) × Golden Delicious (S2S3) 1, 2, 5 USA
6 Gamhong (S1S9) Spur Earliblaze (S1S19) × Spur Golden Delicious (S2S3) 4 Korea
7 Green Ball (S3S7) Golden Delicious (S2S3) × Fuji (S1S9) 1, 2, 5 Korea
8 Hongan (S1S3) Fuji (S1S9) × Jonathan (S7S9) 1, 2, 5 Korea
9 Honggeum (S3S7) Senshu (S1S7) × Hongro (S1S3) 1, 2, 4, 5 Korea
10 Hongro (S1S3) Spur Earliblaze (S1S19) × Spur Golden Delicious (S2S3) 4 Korea
11 Hongso (S1S3) Yoko (S3S9) × Hongro (S1S3) 3, 4, 5 Korea
12 Hwahong (S3S9) Fuji (S1S9) × Sekaiichi (S3S9) 1, 5 Korea
13 Hwangok (S3S9) Kogetsu (S3S7) × Yataka Fuji (S1S9) 1 Korea
14 Manbok (S1S9) Earliblaze (S1S19) × Fuji (S1S9) 1, 4, 5 Korea
15 Picnic (S1S5) Fuji (S1S9) × Sansa (S5S7) 1, 4, 5 Korea
16 Sansa (S5S7) Gala (S2S5) × Akane (S7S24) 1, 4, 5 Japan
17 Seohong (S3S9) Tsugaru (S3S7) × Chukwang (S3S9) 1, 2, 3, 4, 5 Korea
18 Sinano Sweet (S1S7) Fuji (S1S9) × Tsugaru (S3S7) 1, 2, 3, 5 Japan
19 Summer Dream (S7S9) Tsugaru (S3S7) × Natsumidori (S3S9) 1, 2, 3, 5 Korea
20 Summer King (S3S9) Fuji (S1S9) × Golden Delicious (S2S3) 1, 2, 5 Korea
21 Sunhong (S3S9) Hongro (S1S3) × Chukwang (S3S9) 3, 4 Korea
22 Tsugaru (S3S7) Golden Delicious (S2S3) × Unknown 1, 2, 5 Japan
23 Yeohong (S1S9) Jonathan (S7S9) × Fuji (S1S9) 1, 2, 5 Korea
24 Yoko (S3S9) Golden Deliciou s(S2S3) × Unknown 1, 2, 5 Japan
Crabapple
25 Golden Bell (S2S23) Gala (S2S5) × Mo.84 (S4S26) 1, 2, 5 Korea
26 Malus prunifolia (S7S29) Wild species China
27 Maypole (S10S16) McIntosh Wijcik (S10S25) × Malus Baskatong (S16S26) 1, 2 France
28 Purple Lemoine (S7S9) Malus × atrosanguinea × Malus niedzwetskyana - France
29 Rose Bell (S21S26) Gala (S2S5) × Mo.84 (S4S26) 1, 2, 5 Korea
30 Sentinel (S21S26) Unknown - New Zealand
31 Yantaishagou (S9S21) Unknown - -

zParentage’s S-genetypes were reported by references. 1, Broothaerts (2004); 2, Heged s (2006); 3, Heo et al. (2011); 4, Kim et al. (2006); 5, Yamane & Tao (2009)

S-allele specific primers used for PCR amplification.

S-allele Primer Sequence (5’→3’) Amplicon size (bp) Annealing temp. (°C) Reference
S1 MdS1SpF TGTAAGGCACCGCCATATCATAC 734 62 Long et al. (2010)
MdS1SpR CAACCTCAACCAATTCAGTCAATGA
S2 MdS2SpF AACATGAATCGAAGTGAATTATTTA 489 55 ”
MdS2SpR TTGAGGTTTGGTTCCTTACCATG
S3 MdS3SpF GGCGAAAATTAAACCGGAGAAGAA 292 58 ”
MdS3SpR CCTCTCGTCCTATATATGGAAATCAC
S4 MdS4SpF ATTGCAAGACAAGGAATCGTCGGAA 533 63 ”
MdS4SpR AGAAATGTGCTCTGTTTTTATCG
S5 FTC10 CAAACATGGCACCTGTGGGTCTCC 346 60 Broothaerts (2003)
FTC11 TAATAATGGATATCATTGGTAGG
S7 MdS7SpF AGTAAATCAACCGTGGATGCTCAG 397 63 Long et al. (2010)
MdS7SpR TTACAATATCTACCTGTTTCCTGGG
S9 MdS9SpF CCACTTTAATCCTACTCCTTGTAGA 522 63 ”
MdS9SpR TCAATTTCCTTCTGTGTCCTGAATT
S10 FTC12 CCAAACGTACTCAATCGAAG 209 66 Broothaerts (2003)
FTC228 ATGTCGTCCCGTGTCCTGAATC
S11 MdS11SpF AAATATTGCAAGGCGCCGC 678 63 Long et al. (2010)
MdS11SpR TTTCAATATCTACCAGTCTCCGGC
S16 FTC5 TCCCACAATACAGAACGAGA 274, 243+31 (TaqI) 60 Broothaerts (2003)
OWB249 CAATCTATGAAATGTGCTCTG
S19 MdS19SpF GCCTTCAAACAAGAATGGACC 481 63 Long et al. (2010)
MdS19SpR TCAATATCCACCAATGACCTGTT
S20 MdS20SpF GTTGTGGCCTTCAGACTCG 882 64 Long et al. (2010)
MdS20SpR GGCCAACTACTTTTATTTTTCATC
S21 MdS21SpF AAGTAATTGCCCGATAAGGAACATA 584 63 Long et al. (2010)
MdS21SpR AGTTTATGAAATGTTCTCCGCTGTA
S23 FTC222 CAATCGAACCAATCATTTGGT 237 60 Broothaerts (2003)
FTC224 GGTGTCATATTGTTGGTACTAATG
S24 MdS24SpF ATGGCTCCTGTGCGTCTTCCC 421 61 Long et al. (2010)
MdS24SpR CGTCATCCGTGTATAGGGCAACT
S26 MdS26SpF TCCATCAAACGTGACTTCTCAT 423 55 ”
MdS26SpR ATCCTTCAGCATCCTGATTCG
S29 A29PL TATAAACATGGCTCCTGTGCG 217 60 Son et al. (2013)
A29PR TCGTTTGGCACTTGAGTTTTG
S31 MdS31SpF TGACCCAAAATATTGCAAGGCGC 556 63 Long et al. (2010)
MdS31SpR TTTCAATATCTACCAGTCTCCGGC
S44 MdS44SpF GCATGGTAGGACCTGACCCAAGTA 561 63 ”
MdS44SpR TCTCAACCAATTGAGTCGTCGTACC
S45 MdS45SpF CCAGAAGGTTGCAAGACACAGAAAT 641 63 ”
MdS45SpR AGTTTTGGTGCCTTATTGTTGGTAC
S46 MdS46SpF CCCAACGTGCTCGATCGAACA 212 61 ”
MdS46SpR TCAATTTCCTTCTGTGTCCAGAATC

S-allele genotypes of apple and crabapple cultivars by S-allele-specific PCR.

S-allele genotype Cultivar
S1S3 Hongan, Hongro, Hongso
S1S5 Picnic
S1S7 Sinano Sweet
S1S9 Alps Otome, Fuji, Gamhong, Manbok, Yeohong
S2S5 Gala
S2S23 Golden Bell
S3S7 Arisoo, Green Ball, Honggeum, Tsugaru
S3S9 Chukwang, Hwahong, Hwangok, Seohong, Summer King, Sunhong, Yoko
S5S7 Sansa
S7S9 Summer Dream, Purple Lemoine
S7S29 Malus prunifolia
S9S21 Yantaishagou
S10S16 Maypole
S21S26 Rose Bell, Sentinel
Table 1. List of apple and crabapple cultivars used in this study.

Parentage’s S-genetypes were reported by references. 1, Broothaerts (2004); 2, Heged s (2006); 3, Heo et al. (2011); 4, Kim et al. (2006); 5, Yamane & Tao (2009)

Table 2. S-allele specific primers used for PCR amplification.
Table 3. S-allele genotypes of apple and crabapple cultivars by S-allele-specific PCR.

Table 1.

Table 2.

Table 3.

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