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

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개화시기가 상이한 두 개의 무 계통간 개화유전자 서열변이 분석

Sequence Variation Analysis of Flowering-time Genes in Two Radish Lines with Different Flowering Time

Korean Journal of Breeding Science 2017;49(4):351-358.
Published online: November 30, 2017

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*Corresponding author : (yskim0907@naver.com), +82-31-8053-8400, +82-31-8053-8409
*Corresponding author : (hscho@mail.kribb.re.kr), +82-42-860-4469, +82-42-861-1759
• Received: August 23, 2017   • Accepted: August 31, 2017

© Korean Society of Breeding Science. All rights reserved.

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  • It is important for radish to have late flowering characteristics especially in the case of spring and winter cultivars. To understand late flowering characteristics of radish at the molecular level the flowering time genes of two radish lines (NH-JS1 and NH-JS2) with different flowering time were compared by re-sequencing their genomes. There were a total of 872,587 SNPs and 194,637 INDELs between the two lines. The SNP density of each chromosome was relatively uniform throughout, but the region with low SNP density was found at the end of R3 and the middle of R9. To compare the flowering time genes of the two lines, we first looked for the flowering time genes in radish using Arabidopsis thaliana flowering time genes. As a result, homologs of radish were found for most flowering time genes, but FRIGIDA was not found. Among 224 radish flowering time gene-homologs found, 97 genes showed more than one sequence difference (SNP or INDEL) between the two lines, and 127 genes had no difference. In particular, no sequence differences were found in FT, CO, and FLC, core flowering time control genes. Rs350520 (FVE), Rs193800 (CURLY LEAF) and Rs255320 (ATX1) with more than 100 sequence variations were expected to have a significant effect on flowering time difference between the two lines. These results will be of great help in understanding the flowering timing difference between the two lines at the molecular level.
무(Raphanus sativus L.)는 십자화과에 속하는 초본식물로 전세계적으로 식용으로 많이 사용되고 있다. 두꺼운 뿌리뿐 만 아니라 잎도 요리 재료로 사용되고 있으며 특히 국내에서는 김치재료로 사용되어 주요 채소작물로 볼 수 있다. 국내 종자매출 로 보면 고추에 이어 두 번째로 큰 시장을 형성하고 있다.
무의 게놈은 약 570Mb로 추정되며 그 서열은 2014년 처음 발표되었으나 유전자 구조나 발현 연구에 충분하지 않아서 2015 년에 보강된 서열이 발표되었다(Kitashiba et al. 2014, Mitsui et al. 2015). 2016년에는 Jeong 등이 좀 더 세밀한 형태의 서열결 과를 발표하였다 (Jeong et al. 2016). 예상되는 유전자 수는 연구그룹에 따라 다르나 약 46,000~64,000개 일 것으로 추정되 고 있다(Mitsui et al. 2015, Jeong et al. 2016, Li et al. 2016).
무는 작형에 따라 봄무, 여름무, 가을무, 월동무 등으로 나뉘어 진다. 그 중 봄무와 월동무의 경우 낮은 온도에 노출될 가능성이 높고 저온에 노출된 무는 일찍 개화함으로써 무의 상품성을 떨어뜨린다. 따라서 봄무와 월동무의 경우 만추대 특성을 가지고 있는 것이 중요하다. 무 개화에 대한 분자생물학적 연구는 비교적 최근 들어 연구되기 시작하였다(Park et al. 2015, Jung et al. 2016, Li et al. 2016, Nie et al. 2016a, 2016b, Liu et al. 2017). 대표적인 개화촉진 유전자인 FLOWERING LOCUS T (FT), CONSTANS (CO) 등은 무가 생식생장으로 들어서면서 발현 이 증가하나 개화억제 유전자인 FLOWERING LOCUS C (FLC) 등은 발현이 감소하였다(Nie et al. 2016a). 또한 춘화처리 시 FLC 발현양은 감소하는 반면 VERNALIZATION INSENSITIVE 3 (VIN3), VERNALIZATION 1 (VRN1) 등은 발현이 증가하여 애기장대와 상당히 유사하였다 (Jung et al. 2016).
만추대 특성을 가진 계통을 육성하기 위해서는 무의 개화 조절 메커니즘을 이해하는 것과 더불어 만추대 계통과 조추대 계통의 차이를 분자생물학적으로 이해하는 것이 필요하다. 저온 처리 시 만추대 계통은 개화가 빠른 조추대 계통에 비해 개화촉진 유전자의 발현이 낮은 것으로 밝혀졌다(Jung et al. 2016). 특히 SOC1의 발현양이 크게 차이 났으며 그 외 춘화처리에 관련된 유전자의 발현도 의미 있게 차이가 났다. 반면 FLC와 같은 개화 억제 유전자의 발현양은 오히려 더 높아 두 계통간 개화특성 차이를 잘 설명해 주었다. 만추대 계통은 조추대 계통과 비교했을 때 다수의 개화관련 유전자발현 수준이 다른 것이 밝혀졌고 이는 만추대 계통을 육성하는 것이 어려운 이유를 분자생물학적 으로 설명해 주는 결과였다. 본 연구는 이러한 만추대 계통과 조추대 계통의 차이를 보다 세밀하게 연구하기 위해 두 계통을 re-sequencing 하여 게놈 수준에서 개화유전자 차이를 분석하였 다. Blast를 통하여 애기장대 개화 유전자와 비교하여 무 개화 유전자를 조사하였으며 두 계통간 개화 유전자 서열을 비교 하였다.
실험 재료
농협종묘센터에서 보유한 13개 계통에 대해 개화시기를 조사 한 후 개화시기가 많이 차이 나는 만추대 1계통(NH-JS1)과 조추대 1계통(NH-JS2)을 선발하여 실험 재료로 사용하였다 (Park et al. 2015).
Re-sequencing
Re-sequencing을 수행하기 위해 무 잎에서 genomic DNA extraction kit(바이오니아, 한국)를 사용하여 genomic DNA를 추출하였다. 튜브에 잎 조각과 bead를 같이 넣은 뒤 액체질소로 얼렸다. Tissue Lyzer II(QIAGEN, Germany)로 2분 동안 상온 에서 조직을 파쇄한 후 kit내 사용방법에 따라 genomic DNA를 추출하였다. 추출된 DNA는 아가로스겔 전기영동으로 분해여부 를 확인하였다. Re-sequencing은 테라젠바이오(수원, 한국)에 의뢰하여 진행하였으며 일차적인 SNP및 INDEL 추출 또한 테라젠바이오에서 수행되었다. 얻어진 SNP, INDEL 중 두 계통 에서 동형접합성인 것과 depth가 10 이상인 것들만 선발하여 이후 분석에 사용하였다.
무 개화유전자 분석
무 개화유전자를 찾기 위해 애기장대 유전자를 활용하였다. 애기장대 개화유전자 리스트는 독일 막스플랑크 연구소의 Geor ge Coupland 박사팀의 리스트 (http://mpipz.mpg.de/14637/Ar abidopsis_flowering_genes)를 사용하였다. 그 외 PubMed 에 서 조사를 통해 애기장대 유전자를 더 확보하였다. 애기장대 유전자 서열은 TAIR 데이터 베이스(http://www.arabidopsis.or g/index.jsp)에서 확보하였다. 무 개화유전자를 찾기 위해 애기 장대 유전자 서열을 무 CDS 데이터 베이스(http://radish-genom e.org)를 대상으로 blast를 수행하였다. Blast는 local blast(ftp:// ftp.ncbi.nlm.nih.gov/blast/executables/blast+/LATEST/ )를 자체 서버에 설치하여 사용하였다. Blast 결과를 분석하여 애기 장대 유전자와 크기가 비슷하고 identity가 65% 이상, E값이 1e-25이하인 유전자를 선발하였다.
두 계통간 개화유전자 서열차이는 R에서 스크립트를 작성 하여 re-sequencing 데이터에서 추출하였다. 무 염색체별 SNP 밀도는 1 kb 당 SNP 혹은 INDEL의 수를 re-sequencing 데이 터에서 조사하였고 그 결과를 R을 이용하여 나타내었다.
Re-sequencing
무 계통인 NH-JS1과 NH-JS2는 개화시기가 다르다. 25일간 4도씨에서 춘화처리 후 개화시기를 조사해 보면 NH-JS2는 22일 째 추대를 하는 반면 NH-JS1은 전혀 추대를 하지 않았다(Park et al. 2015). 이러한 형질의 차이를 DNA 수준에서 조사해 보기 위해 두 계통에 대해 re-sequencing을 수행하였다. NH-JS1은 총 12.3 Gb, NH-JS2는 8.6 Gb 정도 서열정보가 얻어졌으며 총 맵핑율은 54%(NH-JS1)와 80%(NH-JS2) 이었다(Table 1). Mapping depth는 각각 16.7X와 19.2X 였으며clean reads의 비율은 NH-JS1은 87.54%, NH-JS2는 90.64%였다. 얻어진 서 열 결과를 이용하여 얻은 NH-JS1과 NH-JS2간 SNP는 총 872,587개였으며 INDEL은 194,637개였다. 염색체별로 보면 SNP는 4만~15만개, INDEL은 1만개~3.5만개로 염색체간 차이가 상당히 컸다(Fig. 1). SNP, INDEL 모두 R3, R7 보다 R4, R5, R6에서 많았다. 이러한 차이는 염색체 크기와 상관성이 있었다. R3(29.1 Mb)와 R7(27.2 Mb)은 비교적 크기가 작은 반면 R4~R6(45.9~53.6 Mb)는 크기가 상대적으로 큰 편이었다. INDEL의 크기를 조사해 본 결과 대부분 2bp이하였으나 20 bp이상인 경우도 853개가 확인되었다. 제일 큰 INDEL은 42 bp 차이가 났다(data not shown).
Table 1
Summary of re-sequencing data from the two radish lines.
Table 1
Sample name Total sequenced reads Mapped reads Mapping rate (%) Properly paired reads Properly paired read rate (%) Mapping depth (x)

NH-JS1 125,474,936 50,912,880 51 35,254,463 28.6 16.7
NH-JS2 88,805,772 58,578,707 83 36,230,548 42.1 19.2
Fig. 1
Number of sequence variations (SNP and INDEL) in each chromosome. The size of each chromosome (Mb) is shown below the SNP data.
KJBS-49-351_F1.gif
NH-JS1과 NH-JS2 간의 차이를 좀 더 상세하게 살펴보기 위해 각 염색체별로 SNP 밀도를 조사해 보았다. 평균적으로 1 kb 당 약 2.5개의 SNP가 발견되고 INDEL은 1 kb당 0.56개가 발견되었다. 차이가 나는 지역이 비교적 골고루 분포하고 있으나 차이가 상대적으로 적은 지역도 발견할 수 있었다(Fig. 2). 그 중에서도 특히 R3 끝부분과 R9 중간 부분에서 SNP 밀도가 낮은 지역이 상당히 크게 자리잡고 있었다. 이러한 현상은 수박에 서도 발견되었다. 8개의 수박 계통을 re-sequencing하여 계통간 서열을 비교한 결과 10번, 11번 염색체에서 상당히 넓은 지역에 서 서열의 차이가 적은 지역이 발견되었다(Kim et al. 2016). 이러한 보존 지역이 생긴 이유는 확실하지 않으나 이 지역의 SNP나 INDEL은 품종판별용 마커로 사용하기에 적합하지 않다 는 것은 알 수 있다. 인간에 의한 선발과정 동안 지속적으로 보존된 지역일 가능성도 있으며 이는 다른 무 계통에서 re-sequencing을 통해 확인 가능할 것이다.
Fig. 2
SNP density map. SNP density was measured as the number of SNP per 1 kb. X axis shows position on chromosomes (kb). Y axis shows the number of SNP in each position. The red boxes show the conserved region between NH-JS1 and NH-JS2 line.
KJBS-49-351_F2.gif
무 개화 유전자
무의 개화유전자를 찾아보기 위해 애기장대 개화유전자를 사용하였다. 독일 막스플랑크 연구소의 George Coupland 박사 의 리스트에서 애기장대 개화 유전자리스트를 확보하였다. 이 리스트에 있는 miRNA및 일부 유전자를 제외하고 총 131개의 개화유전자를 찾을 수 있었다. 찾아진 각 유전자를 무 CDS에 blast를 수행하여 무 homolog를 찾아보았다(Suppl. 1). 그 결과 중 일부를 Table 2에 정리하였다. 개화 조절 핵심유전자 중 일부는 무에서 하나만 있는 경우도 있었으나 일부는 여러 개의 유사한 유전자를 찾을 수 있었다. 이렇게 해서 찾은 무 개화유전 자는 총 224개였다. SUPPRESOR OF OVEREXPRESSION OF CONSTANS1 (SOC1)의 경우 4개의 유사한 유전자를 찾을 수 있었고 CO는 2개, FT는 한 개의 유사한 유전자를 찾았다(Table 2). 무는 저온처리 시 개화가 촉진되는데 이와 관련된 FLC의 경우 3개가 발견되었다. 이는 기존에 Yi 등이 무에서 3개의 FLC 유전자를 찾은 결과와 일치한다(Yi et al. 2014). FRIGIDA(FRI) 는 chromatin modification factor를 통해서FLC 유전자 발현을 조절한다(Choi et al. 2011). 그러나 특이하게도FRI의 경우는 homolog라고 생각되는 유전자가 하나도 발견되지 않았다. Blast 결과를 보면 FRI와 서열 유사성이 있는 유전자가 4개 정도 있는 것으로 보이나 실제 그 결과를 상세히 들여다 보면 유사한 유전자 라고 하기는 어려운 것으로 판단되었다. 가장 E값이 낮은 결과를 보인 유전자는 Rs467270(1281 bp)로 FRI (945 bp)보다 크기가 약 300 bp 정도 더 길고 실제 유사성이 있는 부분은 이 유전자의 앞 쪽 448 bp에서만 확인되었다. FRI쪽에서 보면 FRI의 뒤쪽 450bp에서만 유사성이 보였다. 두 번째로 높은 유사성을 보인 유전자(Rs467280, 999 bp)는 FRI와 크기가 유사하였다. 그러나 유사성을 보인 부분은 앞쪽 약 500 bp 부분으로 뒤쪽 절반은 유사성이 없어 이 유전자도 FRI homolog라고 결론 내리기 어려 웠다. 나머지 두 개의 결과도 비슷하였다. 다만 이들 무 유전자들 은 다 염기서열을 기반으로 예측된 것으로 유전자예측 프로그램 의 오류가 있을 가능성도 배제할 수 없다. 따라서 무에서 FRI homolog가 없다고 단정적으로 말하기는 어렵고 실험적으로 확인해 볼 필요가 있다고 생각된다. 또 다른 가능성으로 무에서는 FRI 대신 다른 유전자가 FLC 유전자를 조절할 수도 있다고 생각된다.
Table 2
Blast results of Arabidopsis flowering time genes against radish genes.
Table 2
Query (Arabidopsis) Length (bp) Sbjct (Radish) No. of diversity (SNP/INDEL) between JS1 and JS2 Sbjct Chr Sbjct Length Identity (%) E Value

AGL18 1152 Rs375720 41 R7 777 86 0
Rs332940 0 R6 777 85 0
Rs262750 24 R5 780 84 0

SOC1 645 Rs364220 0 R6 642 93 0
Rs364160 0 R6 642 93 0
Rs113920 84 R3 642 93 0
Rs210580 0 R4 642 93 0

AGL24 663 Rs056630 0 R2 666 89 0

AP1 771 Rs097510 4 R2 585 93 0
Rs416700 0 R8 579 79 6.00E-141
Rs416660 0 R8 579 79 6.00E-141

CO 1122 Rs398510 0 R7 1128 81 0
Rs398520 0 R7 1065 70 1.00E-152

ELF3 2088 Rs196270 66 R4 1815 76 0
Rs346270 0 R6 2049 72 0

EMF1 3291 Rs066730 29 R2 2997 74 0

EMF2 1896 Rs465100 38 R9 1839 92 0
Rs534260 0 R2 1890 87 0

ESD4 1470 Rs164100 4 R4 1347 87 0
Rs165500 5 R4 1335 87 0

FCA 1518 Rs061670 0 R2 2406 84 0

FLC 591 Rs394120 0 R7 594 88 0
Rs144350 0 R3 594 86 0
Rs067530 0 R2 663 74 0

FPA 2577 Rs361990 0 R6 2556 83 0
Rs264090 0 R5 2610 69 0

FPF1 333 Rs168700 13 R4 333 92 1.00E-133
Rs487780 2 R9 327 90 3.00E-127
Rs394480 0 R7 330 79 2.00E-74

FT 528 Rs094390 0 R2 528 88 0

FVE 1524 Rs326180 80 R6 1521 91 0
Rs350520 117 R6 1488 90 0
Rs483490 0 R9 1494 88 0
Rs048390 73 R2 1458 80 0
Rs483480 0 R9 1002 94 0

FY 1944 Rs142690 0 R3 1977 90 0

GI 3522 Rs018220 0 R1 3537 88 0

LFY 1263 Not found

LHY 1938 Rs000160 0 R1 1950 81 0

MAF1,2,3,5 549~618 Rs309680 0 R6 504 86 4.00E-148
Rs309690 0 R6 600 80 5.00E-102
Rs309700 1 R6 570 77 2.00E-120

VIN3 2221 Rs054070 0 R2 2193 81 0
Rs135740 0 R3 1650 83 0

VRN2 1708 Rs433400 0 R8 1281 83 0

VRN5 1989 Rs241200 47 R5 1806 80 0

VRN1 1519 Rs302600 0 R6 1032 89 0
Rs234830 42 R5 1020 86 0
CAL 1033 Not found

TSF 799 Rs094390 0 R2 528 82 1.00E-149

AGL19 691 Rs429750 50 R8 666 86 0

FRI 945 Not found

AGL17 684 Rs041710 71 R2 687 80 8.00E-171

FD 858 Rs150720 20 R4 912 76 1.00E-156
Rs042390 0 R2 684 79 7.00E-135

FLD 2655 Rs287070 0 R5 2685 85 0

FLK 1734 Rs288110 0 R5 1917 84 0

FRL1 1086 Rs015540 9 R1 1071 88 0
Rs015550 6 R1 1071 87 0
Rs101650 0 R2 1083 80 0

FUL 729 Rs311800 40 R6 717 92 0

FWA 2061 Rs472990 1 R9 2070 68 8.00E-50

GA2ox1 990 Rs570400 10 R2 963 85 0
Rs170920 0 R4 984 84 0

GA2ox2 1026 Rs023230 8 R1 1020 88 0
Rs419290 0 R8 1023 86 0

GA2ox3 1008 Rs357280 82 R6 981 85 0

GA20ox2 1137 Rs092510 0 R2 1140 87 0
Rs473380 17 R9 1143 87 0

GAI 1602 Rs179530 6 R4 1599 83 0

LD 2862 Rs088510 0 R2 2814 85 0

RGA 1764 Rs493610 0 R9 1758 78 0

SMZ 1041 Rs260480 92 R5 1029 84 0

TEM2 1059 Rs097160 0 R2 1029 79 0
Rs416970 19 R8 1065 75 0
두 계통간 개화유전자 차이
NH-JS1과 NH-JS2 계통간 개화 유전자 차이를 알아보기 위해 각 개화유전자에서 두 계통의 서열차이를 조사해 보았다. 조사한 모든 개화유전자에서 발견되는 SNP 혹은 INDEL의 수를 Supplementary Table 1에 정리하였다. 또한 일부 유전자의 데이터를 Table 2에 나타내었다. 주요 개화 조절 유전자인 FT, CO에서는 서열차이가 발견되지 않았다. 또한 SOC1은 네 개의 homolog가 찾아졌는데 한 유전자(Rs113920)에서만 서열차이 가 확인되었다(Table 2, Fig. 3). 이러한 차이는 exon에서만 보이는 것이 아니고 intron에서도 나타나 유전자 전체적으로 차이를 보이는 것을 알 수 있었다(Fig. 3). NH-JS1과 NH-JS2는 춘화처리 후 개화시기 차이가 많이 나타나 춘화처리 관련 유전자 (FLC, VRN1, VIN3 등)의 서열차이도 조사하였다. VRN1은 두 개의 유전자가 있는데 그 중 한 개(Rs234830)에서만 SNP가 발견되었다. VERNALIZATION5(VRN5)는 하나의 homolog가 있고 이 유전자(Rs241200)에서 SNP가 다수 발견되었다(Fig. 3). 그러나 FLC에서는 계통간 서열차이를 전혀 확인할 수 없었 다. 한편 FLC발현을 조사한 결과를 보면 두 계통에서 발현차이가 확인 되었다(Jung et al. 2016). 또한 서열차이가 전혀 없었던 SOC1유전자에서도 계통간 발현차이가 많이 나는 것을 알 수 있었다(Jung et al. 2016). 이러한 발현차이는 프로모터 서열차이 때문에 나타날 가능성이 있으므로 두 계통간 서열차이가 없었던 무 SOC1, FLC homolog의 프로모터에서 SNP/INDEL의 수를 조사해 보았다. 그 결과 프로모터에서는 서열차이가 없거나 혹은 4개가 발견되어 발현차이는 프로모터의 서열 차이보다는 유전자 발현을 조절하는 상위 전사인자의 차이 때문일 것으로 추정된다.
Fig. 3
Genomic structure and position of SNP/INDEL in each gene. The vertical lines indicate the position of SNP/INDELs. The black boxes and intervening lines indicate exons and introns, respectively.
KJBS-49-351_F3.gif
본 연구로 찾은 무 개화유전자 224개 중 두 계통간 서열차이 (SNP 혹은 INDEL)이 한 개 이상 발견되는 유전자는 97개였으 며 전혀 차이가 없었던 유전자는 127개였다. 특히 서열차이가 20개 이상 나는 유전자는 43개로 확인되었다. 100개이상 차이 가 나는 유전자는 3개였으며 이들은 각각 Rs350520(FVE), Rs193800(CURLY LEAF), Rs255320(ATX1)이었다. 이 유전 자들은 두 계통간 개화시기 차이에 많은 영향을 줄 것으로 예상된다.
SNP와 유전자 발현
유전자발현 수준이 유전자 서열 등에 따라 달라질 수 있다는 것을 고려하면 SNP와 유전자 발현 간의 상당한 연관성이 예상된 다. 이를 살펴보기 위해 Jung 등(2016)이 발표한 NH-JS1과 NH-JS2에서의 유전자 발현 수준 비교 결과를 활용하여 두 계통 의 개화 유전자 발현 비율과 SNP 수와의 연관성을 조사해 보았 다. 그 결과 사실상 관련이 없는 것으로 나왔다(Fig. 4). 오히려 상당수의 유전자에서 발현차이는 보이나 SNP는 전혀 발견되지 않았고 발현차이는 없으나 SNP는 많이 발견되는 경우도 많았다. 인간 leucocyte에서 연구한 결과를 보면 약 25만개의 SNP 중 유전자 발현과 상관성이 있는 것은 약 2,300개 정도였다(Heap et al. 2009). 따라서 대다수의 SNP는 유전자 발현과 관련이 없었으며 이는 본 연구에서 얻어진 결과와 유사하였다.
Fig. 4
Relationship between gene expression ratio (NH-JS1 versus NH-JS2) and number of SNP. Expression ratio of flowering time genes in two radish lines (NH-JS1 and NH-JS2) was obtained from Jung et al. (2016).
KJBS-49-351_F4.gif
Yi등은 9개의 무 계통을 대상으로 개화시기를 조사하고 각 계통의 FLC 서열의 차이를 살펴보았으나 SNP 수도 적었고 개화시기와 관련된 것을 찾을 수 없었다(Yi et al. 2014). 그러나 FLC1FLC3의 발현은 개화시기와 관련성을 찾을 수 있었다. NH-JS1과 NH-JS2에서도 FLC 유전자 내에서 SNP가 발견되지 않지만 발현차이는 확인되었다(Park et al. 2015). 이는 FLC 상위 조절 유전자 활성이나 발현양의 차이로 인한 것으로 추정된다.
무는 저온처리 시 개화가 촉진되는 작물로 봄과 겨울에 재배되 는 품종의 경우 만추대 특성을 가지는 것이 중요하다. 무 만추대 특성을 연구하기 위해 개화시기가 상이한 두 무 계통(NH-JS1과 NH-JS2)을 선택하여 re-sequencing을 수행하였고 두 계통간 개화유전자 서열차이를 분석하였다. 두 계통간 SNP는 총 872,587개였으며 INDEL은 194,637개였다. 각 염색체별로 SNP 밀도를 조사해 본 결과 전체적으로 비교적 골고루 분포하고 있으나 R3 끝부분과 R9 중간 부분에서 SNP 밀도가 낮은 지역이 상당히 크게 자리잡고 있었다. 두 계통의 개화유전자를 비교하기 위해 무 개화유전자를 먼저 찾아보았다. 무 개화유전자는 애기장 대 개화유전자를 이용하여 조사하였으며 그 결과 대부분의 개화 유전자에 대해 무의 homolog를 찾을 수 있었으나 FRIGIDA는 찾을 수 없었다. 찾아진 무 개화유전자 224개 중 두 계통간 서열차이(SNP 혹은 INDEL)가 한 개 이상인 유전자는 97개였으 며 전혀 차이가 없었던 유전자는 127개였다. 특히 핵심 개화조절 유전자인 FT, CO, FLC에서는 서열차이가 발견되지 않았다. Rs35020 (FVE), Rs193800 (CURLY LEAF), Rs255320 (ATX1) 는 서열차이가 100개 이상으로 두 계통간 개화시기 차이에 많은 영향을 줄 것으로 예상되었다. 이러한 연구결과는 두 계통간 개화시기 차이를 분자수준에서 이해하는 데 큰 도움을 줄 것으로 기대된다.
본 연구는 농림수산식품부의 재원으로 농림식품기술기회평가 원의 농생명산업기술개발 사업(과제번호: 114061-03-3-HD020 와 116091-03-2-SB010)의 지원으로 수행되었음.
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  • 2. Heap G.A., Trynka G., Jasen R.C., Bruinenberg M., Swertz M.A., Dinesen L.C., Hunt K.A., Wijmenga C., vanHeel D.A., et al. Complex nature of SNP genotype effects on gene expression in primary human leucocytes. BMC Med. Genet. 2009. 2: 1-13.
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Sequence Variation Analysis of Flowering-time Genes in Two Radish Lines with Different Flowering Time
Korean. J. Breed. Sci.. 2017;49(4):351-358.   Published online December 1, 2017
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Sequence Variation Analysis of Flowering-time Genes in Two Radish Lines with Different Flowering Time
Korean. J. Breed. Sci.. 2017;49(4):351-358.   Published online December 1, 2017
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Sequence Variation Analysis of Flowering-time Genes in Two Radish Lines with Different Flowering Time
Image Image Image Image
Fig. 1 Number of sequence variations (SNP and INDEL) in each chromosome. The size of each chromosome (Mb) is shown below the SNP data.
Fig. 2 SNP density map. SNP density was measured as the number of SNP per 1 kb. X axis shows position on chromosomes (kb). Y axis shows the number of SNP in each position. The red boxes show the conserved region between NH-JS1 and NH-JS2 line.
Fig. 3 Genomic structure and position of SNP/INDEL in each gene. The vertical lines indicate the position of SNP/INDELs. The black boxes and intervening lines indicate exons and introns, respectively.
Fig. 4 Relationship between gene expression ratio (NH-JS1 versus NH-JS2) and number of SNP. Expression ratio of flowering time genes in two radish lines (NH-JS1 and NH-JS2) was obtained from Jung et al. (2016).
Sequence Variation Analysis of Flowering-time Genes in Two Radish Lines with Different Flowering Time

Summary of re-sequencing data from the two radish lines.

Sample name Total sequenced reads Mapped reads Mapping rate (%) Properly paired reads Properly paired read rate (%) Mapping depth (x)

NH-JS1 125,474,936 50,912,880 51 35,254,463 28.6 16.7
NH-JS2 88,805,772 58,578,707 83 36,230,548 42.1 19.2

Blast results of Arabidopsis flowering time genes against radish genes.

Query (Arabidopsis) Length (bp) Sbjct (Radish) No. of diversity (SNP/INDEL) between JS1 and JS2 Sbjct Chr Sbjct Length Identity (%) E Value

AGL18 1152 Rs375720 41 R7 777 86 0
Rs332940 0 R6 777 85 0
Rs262750 24 R5 780 84 0

SOC1 645 Rs364220 0 R6 642 93 0
Rs364160 0 R6 642 93 0
Rs113920 84 R3 642 93 0
Rs210580 0 R4 642 93 0

AGL24 663 Rs056630 0 R2 666 89 0

AP1 771 Rs097510 4 R2 585 93 0
Rs416700 0 R8 579 79 6.00E-141
Rs416660 0 R8 579 79 6.00E-141

CO 1122 Rs398510 0 R7 1128 81 0
Rs398520 0 R7 1065 70 1.00E-152

ELF3 2088 Rs196270 66 R4 1815 76 0
Rs346270 0 R6 2049 72 0

EMF1 3291 Rs066730 29 R2 2997 74 0

EMF2 1896 Rs465100 38 R9 1839 92 0
Rs534260 0 R2 1890 87 0

ESD4 1470 Rs164100 4 R4 1347 87 0
Rs165500 5 R4 1335 87 0

FCA 1518 Rs061670 0 R2 2406 84 0

FLC 591 Rs394120 0 R7 594 88 0
Rs144350 0 R3 594 86 0
Rs067530 0 R2 663 74 0

FPA 2577 Rs361990 0 R6 2556 83 0
Rs264090 0 R5 2610 69 0

FPF1 333 Rs168700 13 R4 333 92 1.00E-133
Rs487780 2 R9 327 90 3.00E-127
Rs394480 0 R7 330 79 2.00E-74

FT 528 Rs094390 0 R2 528 88 0

FVE 1524 Rs326180 80 R6 1521 91 0
Rs350520 117 R6 1488 90 0
Rs483490 0 R9 1494 88 0
Rs048390 73 R2 1458 80 0
Rs483480 0 R9 1002 94 0

FY 1944 Rs142690 0 R3 1977 90 0

GI 3522 Rs018220 0 R1 3537 88 0

LFY 1263 Not found

LHY 1938 Rs000160 0 R1 1950 81 0

MAF1,2,3,5 549~618 Rs309680 0 R6 504 86 4.00E-148
Rs309690 0 R6 600 80 5.00E-102
Rs309700 1 R6 570 77 2.00E-120

VIN3 2221 Rs054070 0 R2 2193 81 0
Rs135740 0 R3 1650 83 0

VRN2 1708 Rs433400 0 R8 1281 83 0

VRN5 1989 Rs241200 47 R5 1806 80 0

VRN1 1519 Rs302600 0 R6 1032 89 0
Rs234830 42 R5 1020 86 0
CAL 1033 Not found

TSF 799 Rs094390 0 R2 528 82 1.00E-149

AGL19 691 Rs429750 50 R8 666 86 0

FRI 945 Not found

AGL17 684 Rs041710 71 R2 687 80 8.00E-171

FD 858 Rs150720 20 R4 912 76 1.00E-156
Rs042390 0 R2 684 79 7.00E-135

FLD 2655 Rs287070 0 R5 2685 85 0

FLK 1734 Rs288110 0 R5 1917 84 0

FRL1 1086 Rs015540 9 R1 1071 88 0
Rs015550 6 R1 1071 87 0
Rs101650 0 R2 1083 80 0

FUL 729 Rs311800 40 R6 717 92 0

FWA 2061 Rs472990 1 R9 2070 68 8.00E-50

GA2ox1 990 Rs570400 10 R2 963 85 0
Rs170920 0 R4 984 84 0

GA2ox2 1026 Rs023230 8 R1 1020 88 0
Rs419290 0 R8 1023 86 0

GA2ox3 1008 Rs357280 82 R6 981 85 0

GA20ox2 1137 Rs092510 0 R2 1140 87 0
Rs473380 17 R9 1143 87 0

GAI 1602 Rs179530 6 R4 1599 83 0

LD 2862 Rs088510 0 R2 2814 85 0

RGA 1764 Rs493610 0 R9 1758 78 0

SMZ 1041 Rs260480 92 R5 1029 84 0

TEM2 1059 Rs097160 0 R2 1029 79 0
Rs416970 19 R8 1065 75 0
Table 1 Summary of re-sequencing data from the two radish lines.
Table 2 Blast results of Arabidopsis flowering time genes against radish genes.