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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers
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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers

Byeong Wan Kim1, Kyu Jin Sa1, Ki Jin Park2, Jong Yeol Park2, Ju Kyong Lee1,*

Published online: February 28, 2015

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*Corresponding author (E-mail: jukyonglee@kangwon.ac.kr, Tel: +82-33-250-6415, Fax: +82-33-259-5558)
• Received: February 14, 2015   • Revised: February 28, 2015   • Accepted: March 2, 2015

© The Korean Society of Breeding Science

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  • In this study, genetic diversity, genetic relationship, population structure and association analysis among 12 colored and non-colored maize lines were investigated using 300 simple sequence repeat (SSR) markers. In genetic diversity analysis, a total of 1,331 alleles were identified at all the loci with an average of 4.44 and a range between 2 and 10 alleles per locus. The major allele frequency varied from 0.25 to 0.92 with an average of 0.48. Among 1,331 alleles, 221 alleles were colored maize specific allele, and 408 alleles were non-colored maize specific allele. While 702 alleles were shared in colored and non-colored maize lines. In addition, colored maize specific alleles were detected in 163 SSR markers. From population structure analysis, the 12 colored and non-colored maize lines were separated with based on the membership probability threshold 0.8, and divided into groups I, II, III and admixed group. The results of this study could serve as useful molecular markers for identification and characterization of colored and non-colored maize lines in the maize breeding studies.
옥수수(Zea mays L.)는 세계 3대 중요 작물 중 하나이며, 식용뿐만 아니라 사료용, 공업용 및 바이오에너지 등 다양한 용도로 이용되고 있다. 최근 웰빙(well-being)에 대한 관심이 높아지면서, 자연적으로 생성되는 천연성분을 함유한 고품질, 고기능성을 갖는 작물이나 이를 이용한 가공품을 선호하고 있다. 더욱이 검은콩, 흑미, 흑깨 등 웰빙 건강식으로 블랙푸 드가 각광을 받으면서 이러한 블랙푸드에 함유된 안토시아닌 (anthocyanin) 색소에 대한 관심이 높아지고 있다. 안토시아 닌 색소는 식물에 존재하는 천연 색소로 자색, 청색, 적색을 나타내며 인체에 무해한 성분이다. 안토시아닌 색소는 천연 착색료(Cevallos-Casals et al. 2004)로 이용될 뿐 아니라, 항 산화(Yoshimoto et al. 2001), 항암(Hagiwara et al. 2001), 항당뇨(Li et al. 2011)에 효과가 있었다. 현재 안토시아닌 색 소는 포도에서 주로 추출되어 이용되고 있다(Kim et al. 2008). 이러한 안토시아닌 색소의 중요성과 기능성 등을 고려 하여, 강원도농업기술원 옥수수연구소에서는 안토시아닌 색소 이외에 유용 기능성 물질을 다량 보유하고 있는 기능성 옥수수 의 품종개발에 관심을 갖고 있다. 하지만 현재까지 색소옥수수 품종의 개발 및 분자생물학적 연구가 매우 제한적이며, 다양한 특성을 갖는 색소옥수수 품종의 개발이 절실히 요구된다.
색소옥수수의 품종개발을 위해서는 우수 형질을 지니고 있 는 다양한 유전자원의 수집과 수집된 유전자원들의 유전학적 특성을 명확히 밝히는 것이 육종의 효율을 높이기 위해 매우 중요하다. 특히 타식성 작물인 옥수수 품종의 경우, 다양한 지 역에서 수집되어 고정된 육종재료들에서 유전자원들의 유전 학적 특성을 밝히고, 우수한 자식계통을 선정하기 위해서 수 집된 재료들의 유전적 다양성(genetic diversity), 계통유연관 계(genetic relationship) 및 집단구조(population structure)를 명확히 평가하는 것이 신품종 개량을 위한 자식계통 육성 및 발굴 그리고 교배조합 예측 등에 효율적으로 이용될 수 있고, 이형집단에서 자식계통의 선발과 품종 보호 등에도 매우 중 요하다(Hallauer et al. 1988, Pejic et al. 1998). 특히 본 연 구에서 분석에 이용한 SSRs (simple sequence repeats) 분석 법은 PCR을 이용하는 분석기법으로 재현성이 매우 뛰어나 며, 식물 집단들 내에서 유전자좌마다 많은 대립인자들을 가 지고 있고 대립유전자의 특성을 명확하게 나타낼 수 있어 옥 수수와 같은 작물의 유전적 다양성, 계통유연관계 및 집단구 조를 분석하고 fingerprinting과 molecular mapping 등에 활 용되고 있다(Hu et al. 2009).
본 연구는 강원도 농업기술원 옥수수연구소에서 고품질의 색소옥수수 품종을 개발하기 위해서 육성한 12개 색소 및 비 색소옥수수 핵심집단에 대하여 SSR 분자마커를 이용하여 유 전적 다양성, 계통유연관계 및 집단구조 분석을 수행하고, 색 소 및 비색소옥수수 그룹에서 특이적으로 나타나는 대립단편 을 선발하여 색소옥수수 육종에서 색소옥수수 계통 선발에 유용한 정보를 제공하고자 하였다.
공시재료 및 DNA 추출
본 연구에 이용된 색소 및 비색소옥수수 핵심집단 계통들 은 강원도 농업기술원 옥수수연구소에서 기능성 색소옥수수 및 종실옥수수 신품종 육성을 위해 육성한 자식계통들로, Kim et al. (2012)이 보고한 31개의 색소 및 비색소옥수수 자 식계통들에 대한 dendrogram 분석 결과에 기초하여 색소 및 비색소옥수수 집단으로 계통유연관계가 명확히 나타나는 12 개의 옥수수 핵심집단(색소옥수수 6계통, 비색소옥수수 6계 통)을 선발하였다(Table 1). 옥수수 genomic DNA는 유묘기 의 어린 잎에서 Dellaporta et al. (1983)의 방법에 따라 추출 하였다.
Table 1.
Derivation of 12 colored and non-colored maize lines used in this study.
Table 1.
Code No. Entry No. Pedigree Type
1 10S4015 China collection 1-1 Colored kernel, normal corn
2 10S4026 China collection 1-4 Colored kernel, normal corn
3 10S4032 China collection 1-7 Colored kernel, normal corn
4 11CS4117 HW9 BC2 Colored kernel, waxy corn
5 11CS4124 HW9 BC2 Colored kernel, waxy corn
6 11CS7014 HW9 BC1 Colored kernel, waxy corn
7 Mo17 Mo17 Normal corn, inbred line
8 B14A B14A Normal corn, inbred line
9 HW7 HW7 Waxy corn, inbred line, black kernel
10 KW7 KW7 Waxy corn, inbred line, black kernel
11 HW9 HW9 Waxy corn, inbred line, black kernel
12 HW3 HW3 Waxy corn, inbred line, black kernel
SSR 분석 및 PCR 증폭
색소 및 비색소옥수수 핵심집단에 대한 유전적 다양성 및 집단구조 분석을 위해서 선발된 SSR primer들은 MaizeGDB (http://www.maizegdb.org/) 정보를 기초하여, 옥수수 전체 게놈을 포함할 수 있도록 각 염색체 당 30개씩, 총 300개의 SSR primer를 선발하였다.
PCR 증폭에서 용액 조성은 총 30 mL로 20 ng의 genomic DNA, 1x PCR buffer, 0.3 mM의 forward와 reverse primers, 0.2 mM dNTPs, 1 unit의 Taq Polymerase (Biotools, B&M Labs, Madrid)로 구성하였다. PCR 반응은 처음 94°C에서 5분간 최초 denaturation을 한 후, 다음 94°C에서 1분간 denaturation, 65°C에서 1분간 annealing, 72°C에서 2분간 extension을 하 였으며, 2번째 cycle부터 annealing 온도를 1°C씩 낮추어 최 종 55°C까지 20 cycle을 반복한 후, final extension을 72°C 에서 10분간 실시하였다.
전기영동 및 silver-staining
PCR 증폭이 끝난 product 5μl를 10 μl의 loading-buffer (98% formamide, 0.02% BPH, 0.02% Xylene C, 5 mM NaOH)와 혼합하였다. 그 후 denaturation 처리를 한 후 얼음에 냉 각하여, 6% denaturing (7.5 M urea) acrylamide-bisacrylamide gel (19:1)에 1x TBE buffer를 처리 후 2 μl의 sample을 로 딩하고, 1,800 V, 60 W에서 2시간 동안 전기영동을 실시하 였다. 전기영동이 끝난 gel은 silver-staining kit (Promega, USA)를 이용하여 DNA band를 확인하였다.
Data 통계 분석
통계분석은 SSR primer들에서 증폭된 DNA 단편을 대립 단편(allele)의 유무에 따라 1(유)과 0(무)으로 기록하였다. Locus당 대립단편 수, major allele frequency (MAF), gene diversity (GD), polymorphic information content (PIC)는 PowerMarker V3.25 프로그램을 이용하여 분석하였다(Liu & Muse 2005). 유전적 다양성 값은 다음과 같이
Gene diversity GD=1u=1kplu2/11fn
의 공식으로 계산하였으며, 여기서 plu는 u번째 allele의 빈도, f는 inbreeding coefficient, n은 sample size를 나타낸다. 그 리고 PIC값은 Bostein et al. (1980)의 계산 방식에 따라 아래 와 같이
PIC=1u=1kplu2u=1k1=u1k2plu2pl2
의 공식을 이용하였으며, 여기서 plu는 u번째의 allele 빈도, plv는 v번째의 allele 빈도를 나타낸다.
유전적 유사성(genetic similarities, GS)값은 Dice similarity index (Dice 1945)를 이용하여 계산하였고, dendrogram (SAHN Clustering)은 NTSYS-pc.V.2.1 프로그램을 이용하여 UPGMA (un-weighted pair group methods using arithmetic averages algorithm)방법에 따라 작성하였다(Rohlf 1998). 한편 색소 및 비색소옥수수 계통들에 대한 집단구조 분석은 model-based program인 STRUCTURE 2.3를 이용하여 Pritchard et al. (2003)가 제안한 방법에 의해 계산하였다. STRUCTURE program을 이용한 분석에서 admixture model은 100,000의 burn-in, 100,000의 run length로 1~10의 범위에서 각 K값 당 5반복을 실시하여 평균 likelihood value인 LnP(D)값을 계산하였으며, STRUCTURE program에서 측정된 데이터의 log probability [LnP(D)]는 정확한 subgroup 수를 평가할 수 없어서, Evanno et al. (2005)가 제안한 ad hoc criterion (Δ K)를 이용하여 K값을 결정하였다.
색소 및 비색소옥수수 핵심집단의 유전적 다양성 및 색소 특이적 마커 선발
본 연구에서는 색소 및 비색소옥수수 12계통(색소옥수수 6 계통, 비색소옥수수 6계통)에 대하여 300개의 SSR primer를 이용하여 색소 및 비색소옥수수 계통들간의 유전적 변이성을 분석하였다. 그 결과 대립단편의 크기가 54 bp에서 396 bp의 범위로 나타났으며, 총 1,331개의 대립단편을 증폭시켰다. SSR primer당 증폭된 대립단편들은 최소 2개에서 최대 10개 까지 나타났으며, SSR primer당 평균 4.44개가 증폭되었다. 그리고 300개의 SSR primer들에서 측정된 MAF는 0.25에서 0.92의 범위로 나타났고, 평균값은 0.48을 나타내었다. 유전 적 다양성(GD)값은 0.15에서 0.88의 범위로 나타났으며, 평 균 0.64의 값을 나타내었다. PIC값은 0.14에서 0.86의 범위 로 나타났으며, 평균 0.59의 값을 나타내었다(Table 2).
Table 2.
Mean of characteristics of 300 SSR loci in 12 colored and non-colored maize lines.
Table 2.
Chr No. of allele MAF* GD** PIC
1 4.20 0.48 0.65 0.59
2 4.83 0.47 0.66 0.61
3 3.90 0.55 0.58 0.53
4 4.83 0.43 0.67 0.62
5 4.37 0.46 0.64 0.59
6 4.73 0.50 0.62 0.58
7 4.37 0.48 0.64 0.60
8 4.40 0.47 0.64 0.59
9 4.47 0.49 0.63 0.58
10 4.27 0.48 0.62 0.57
Total 1,331
Mean 4.44 0.48 0.64 0.59

*MAF: Major Allele Frequency

**GD: Genetic diversity

300개의 SSR primer를 이용하여 색소옥수수 6계통과 비 색소옥수수 6계통의 유전적 변이성을 각각 분석하여 비교를 해보았다. 그 결과 증폭된 총 대립단편의 수는 색소옥수수가 833개, 비색소옥수수가 1,010개로 비색소옥수수가 더 많은 대립단편이 증폭되었고, MAF는 색소옥수수에서 0.57, 비색 소옥수수에서 0.51의 평균값을 보이며 색소옥수수가 더 높은 수치를 나타내었고, 평균 GD 및 PIC값은 색소옥수수 계통에 서는 0.52, 0.45 그리고 비색소옥수수 계통에서는 0.59, 0.54 로 비색소옥수수에서 높은 수치를 나타내었다. 이러한 결과로 보아 두 집단 사이에 큰 차이를 보이지는 않았지만 전체적으 로 비색소옥수수에서 높은 수치를 나타내었다(Table 3).
Table 3.
Mean of number of alleles and genetic diversity index for 300 SSR loci between colored and non-colored maize lines.
Table 3.
Chr Colored maize
 Non-colored maize
No. of Allele MAF* GD** PIC No. of Allele MAF* GD** PIC
1 3.13 0.50 0.59 0.52 3.07 0.54 0.57 0.51
2 3.00 0.55 0.54 0.49 3.30 0.50 0.60 0.54
3 2.77 0.59 0.51 0.44 2.70 0.61 0.49 0.43
4 3.23 0.52 0.57 0.51 3.67 0.46 0.64 0.58
5 2.80 0.57 0.52 0.46 3.10 0.55 0.56 0.50
6 2.53 0.64 0.44 0.39 3.67 0.48 0.63 0.58
7 2.67 0.58 0.52 0.45 3.47 0.49 0.61 0.56
8 2.53 0.60 0.48 0.41 3.67 0.47 0.64 0.58
9 2.37 0.58 0.47 0.40 3.73 0.47 0.64 0.58
10 2.73 0.57 0.51 0.45 3.30 0.52 0.59 0.53
Total 833 1,010
Mean 2.78 0.57 0.52 0.45 3.37 0.51 0.59 0.54

*MAF: Major Allele Frequency

**GD: Genetic diversity

300개의 SSR primer를 이용하여 색소옥수수 그리고 비색 소옥수수 계통들에 대하여 증폭된 1,331개의 대립단편들의 빈도를 확인하였다. 그 결과 증폭된 1,331개의 대립단편들 중 에서 406개(30.5%)의 대립단편들은 private allele를, 566개 (42.5%)의 대립단편들은 rare allele(빈도 <0.1)를, 681개 (51.1%)의 대립단편들은 intermediate allele(빈도 0.1~0.5) 를, 그리고 84개(6.3%)의 대립단편들은 abundant allele(빈도 >0.5)를 각각 나타내었다(Fig. 1). 또한 본 연구에서는 총 12 계통의 옥수수에서 확인된 1,331개의 대립단편 중에서 각 그 룹(색소 및 비색소)별로 특이적으로 나타나는 대립단편을 분 석하였다. 그 결과 총 221개의 대립단편들이 색소옥수수 계 통들에서만 특이적으로 나타났으며, 총 408개의 대립단편은 비색소옥수수 계통들에서만 특이적으로 확인되었다. 더욱이 나머지 702개의 대립단편들은 색소와 비색소옥수수 계통들에서 공통으로 증폭되었다(Fig. 2). 한 예로 SSR primer bnlg1867의 경우 6개의 색소옥수수 자식계통들에서만 특이적으로 2개의 대립단편이 확인되었는데, 그 중 하나는 중국 색소옥수수 3계 통들에서만, 다른 하나는 일반 색소옥수수 3계통들에서만 나 타났다(Fig. 3).
Fig. 1.
Histogram of allele frequencies for the 1,331 alleles in 12 accessions of colored and non-colored maize lines.
KJBS-47-54_F1.gif
Fig. 2.
The Venn diagram of the number of colored and non-colored specific alleles and shared alleles.
KJBS-47-54_F2.gif
Fig. 3.
An example of a SSR profile in 12 colored and non-colored maize lines using the SSR primer, bnlg1867. ★: Specific allele to China colored maize lines, ▲: Specific allele to colored normal maize lines.
KJBS-47-54_F3.gif
색소 및 비색소옥수수 핵심집단의 계통유연관계 및 집단구조
본 연구에서는 12개의 색소 및 비색소옥수수 계통들에 대 한 집단구조와 계통유연관계를 분석하였다. 집단구조 분석을 위해 STRUCTURE program을 이용하여 분석한 결과 ΔK 값이 K=3에서 가장 높은 값이 확인되었다. ΔK 값이 K=3일 때 분석 결과를 보면 Group I, II, III, admixed로 구분되었고, 그 중 Group I은 중국의 색소옥수수 2계통(10S4015, 10S4026) 이며, Group II는 색소종실옥수수 3계통(11CS4117, 11CS4124, 11CS7014)과 찰옥수수 계통인 HW9이며, Group III는 비색 소옥수수 4계통(Mo17, B14A, HW7, HW3)으로 나타났고, admixed group은 찰옥수수 계통인 KW7과 중국 색소옥수수 계통인 10S4032으로 구성되어 있었다(Fig. 4).
Fig. 4.
UPGMA dendrogram and population structure based on the 300 SSR markers in 12 colored and non-colored maize lines. □: colored maize lines; ■: non-colored maize lines.
KJBS-47-54_F4.gif
한편 본 연구에서 색소 및 비색소옥수수 계통들간의 계통유 연관계를 명확히 보기 위해서 UPGMA법에 의한 dendrogram 을 작성하였다(Fig. 4). 그 결과 12계통의 옥수수 계통들은 유 전적 유사성이 30%에서 크게 4개의 Group으로 나누어졌다. Group I은 중국의 색소옥수수 3계통(10S4015, 10S4026, 10S4032)과 종실옥수수 계통인 Mo17을 포함하고 있었고, Group II는 색소종실옥수수 3계통(11CS4117, 11CS4124, 11CS7014)과 찰옥수수 계통인 HW9과 KW7을 포함하였고, Group III는 비색소옥수수 계통인 B14A와 HW3를 포함했다 그리고 Group IV는 찰옥수수 계통인 HW7을 포함했다. Group I의 경우 유전적 유사성 33% 수준에서 2개의 subgroup 으로 나누어 졌다. 첫 번째 subgroup에는 중국의 색소옥수수 3계통(10S4015, 10S4026, 10S4032)이 포함되었고, 두 번째 subgroup에는 종실옥수수인 Mo17이 포함되었다. Group II 의 경우는 유전적 유사성 44.5% 수준에서 2개의 subgroup으 로 나누어졌다. 첫 번째 subgroup에는 색소종실옥수수 3계통 (11CS4117, 11CS4124, 11CS7014)과 찰옥수수 계통인 HW9 이 포함되었고, 두 번째 subgroup은 찰옥수수 계통인 KW7 이 포함되었다(Fig. 4). 이러한 결과를 바탕으로 색소옥수수 핵십집단 3계통(11CS4117, 11CS4124, 11CS7014)과 비색 소옥수수 핵심집단 3계통(Mo17, B14A, HW7)을 선발하여 색소 및 비색소옥수수 집단에서 특이적으로 나타나는 분자마 커를 분석하였다. 그 결과 300개의 SSR primer 중 163개의 primer들이 색소옥수수 집단들에서 특이적으로 나타났다. 색소 특이적 대립단편의 수는 마커 별로 1~3개였으며, bnlg1036, bnlg1662, mmc0111, dupssr24, umc2278, bnlg1046, umc1248, umc1018, umc1002 등의 SSR primer들에서 3개의 색소 특 이적 대립단편이 확인되었다(Table 4).
Table 4.
Identification of colored maize specific markers among 300 SSR loci.
Table 4.
본 연구에서는 강원도농업기술원 옥수수연구소에서 기능 성 색소옥수수 품종개발을 위하여 육성 중인 색소 및 비색소 옥수수 자식계통들에 대하여 유전적 변이성, 집단구조 및 계 통유연관계 분석을 실시하였다. 300개의 SSR primer를 이용 하여 12계통의 색소 및 비색소옥수수 계통들에 대한 유전적 다양성 분석한 결과, 54 bp에서 396 bp의 범위로 나타났으며, 총 1,331개의 대립단편을 증폭시켰고, SSR primer당 증폭된 대립단편들은 최소 2개에서 최대 10개까지 나타났으며, 평균 4.44개가 증폭되었다. MAF, GD, PIC값은 각각 평균 0.48, 0.64, 0.59의 값을 나타내었다(Table 2). 색소옥수수 6계통과 비색소옥수수 6계통의 대립단편 수, GD, PIC를 각각 비교한 결과, 비색소옥수수 계통들이 색소옥수수 계통들보다 높은 값 을 보였다(Table 3). 또한 12계통의 옥수수에서 확인된 1,331 개의 대립단편 중 221개의 대립단편들이 색소옥수수 계통들 에서만 특이적으로 나타나는 것으로 확인되었다.
다양한 계통들에 대하여 Genotype level에서 유전적 다양 성을 평가하는 방법 중에서 가장 단순하고 직접적인 방법은 SSR 마커에서 증폭된 대립단편의 수를 비교하는 것이다 (Cömertpay et al. 2012). 그 결과, 분석에 이용된 SSR primer 들 중에서 dupssr30b, umc1757, bnlg1784, phi026, bnlg1444, bnlg1118, umc1018, bnlg2235, bnlg1525, bnlg1028, bnlg2082, mmc0111 등의 SSR primer들은 색소 및 비색소옥수수 계통 들에서 8~10개의 대립단편 수를 나타냈으며, 반면에 umc1515, umc2249, umc1158, umc1659, phi102228 등의 21개의 SSR primer들은 단지 2개의 대립단편을 나타내었다. 본 연구에서 확인된 많은 대립단편을 나타내는 SSR primer들은 이 후의 옥수수 유전적 다양성 평가에 매우 유용한 마커들로 생각된 다. 더욱이 색소옥수수 계통들에서만 특이적으로 나타난 221 개의 대립단편들은 색소 관련 분자마커를 개발하는데 매우 유용할 것으로 생각되었다. 마지막으로 본 연구에서 색소 및 비색소옥수수 집단에서 측정된 유전적 다양성과 대립단편들 의 빈도 및 SSR primer별 특이적 대립단편들에 대한 정보는 앞으로 강원도농업기술원 옥수수연구소에서 색소옥수수 육종 연구에 유용한 정보를 제공할 것으로 기대되며, 또한 색소와 비색소옥수수 계통들의 식별 및 특정 형질에 연관된 분자마 커 개발에 도움을 줄 것으로 기대한다.
타식성 작물인 옥수수 자식계통들은 육성내력에 따라 genetic background가 매우 복잡하기 때문에 고정된 자식계통들에 대한 계통유연관계 및 집단구조를 이해하는 것이 자식계통 육성 및 보존을 위해서 매우 중요하다(Wang et al. 2008). 본 연구에서 집단구조에 대한 분석결과에서 색소 및 비색소옥수 수 12계통들은 Group I, II, III, admixed로 구분되었고, Group I은 중국의 색소옥수수 2계통, Group II는 색소종실옥 수수 3계통과 찰옥수수 계통인 HW9이 포함되었는데 비색소 옥수수 계통인 HW9이 Group II에 포함된 것은 색소옥수수 계통인 11CS4117, 11CS4124, 11CS7014 자식계통들을 육성 하기 위해 HW9의 여교배로 육성되었기 때문에 같은 Group II 에 속한 것으로 보여진다. Group III는 비색소옥수수 4계통이 포함되었고, admixed group은 찰옥수수 계통인 KW7과 중국 색소옥수수 계통인 10S4032으로 구성되었다. 집단구조분석 결과는 KW7과 중국 색소옥수수 계통인 10S4032을 제외하 고는 전체적으로 기원에 따라 구분되었고, KW7과 HW9을 제외하고는 색소옥수수 계통과 비색소옥수수 계통이 구분되 었다(Fig. 4). UPGMA법에 의한 계통유연관계는 유전적 유 사성이 30%에서 크게 4개의 Group으로 나누어졌다. 비록 색 소옥수수 계통들과 비색소옥수수 계통들이 명확하게 구분되 지 않았지만, 집단구조 분석 결과와 유사하게 중국의 색소옥 수수 3계통이 같은 Group에 포함된 것과 색소종실옥수수 3 계통(11CS4117, 11CS4124, 11CS7014)이 HW9과 같은 Group에 포함되었다. 이러한 결과를 바탕으로 6계통의 색소 및 비색소옥수수 핵심집단을 선발하였고, 300개의 SSR primer 중 핵심집단에서 163개의 primer들이 색소옥수수 집 단들에서 특이적인 마커로 나타났다(Table 4). 6계통의 핵심 집단에 대한 특이적 분자마커들은 앞으로 색소 관련 옥수수 의 육종 연구에 많은 도움이 될 것으로 기대되며, 보다 정확 한 결과를 도출하기 위해서 추가적인 연구가 진행되어야 할 것이다.
본 연구는 색소 및 비색소옥수수 계통들에 대하여 300개의 SSR primer들을 이용하여 유전적 다양성, 집단구조, 계통유 연관계 분석을 수행하였다. 특히 본 연구에서 분석에 이용한 SSR 분자마커는 식물집단 내에서 대립단편의 유무를 쉽게 구분할 수 있는 공우성 마커이므로 12개의 색소 및 비색소옥 수수 집단에 대한 유전적 다양성 및 계통유연관계 그리고 집 단구조를 명확히 나타낼 수 있었다. 따라서 본 연구에서 300 개의 SSR primer들을 이용하여 색소 및 비색소옥수수 계통 들에 대한 분석 결과는 앞으로 강원도농업기술원 옥수수연구 소에서 색소옥수수 자식계통들에 대한 선발과 품종 육성에 유용한 정보를 제공할 것으로 생각된다.
본 연구는 12개의 색소 및 비색소옥수수 계통들에 대하여 300개의 SSR마커를 이용하여 유전적 다양성, 계통유연관계 및 집단구조 분석을 실시하였다. 유전적 다양성 분석 결과, 300개의 SSR primer들은 색소 및 비색소옥수수 12계통들에 서 총 1,331개의 대립단편을 증폭시켰으며, SSR primer당 대 립단편들은 최소 2개에서 최대 10개까지 나타나 평균 4.44개 가 증폭되었다. MAF는 0.25에서 0.92의 범위로 나타났고, 평균값은 0.48을 나타내었다. 총 1,331개의 대립단편 중에서 221개의 대립단편들은 색소옥수수 계통들에서 특이적으로 나 타났고, 408개의 대립단편은 비색소옥수수 계통들에서 특이 적으로 나타났으며, 나머지 702개의 대립단편들은 색소 및 비색소 계통들에서 공통으로 확인되었다. 그리고 총 163개의 SSR 마커에서 색소옥수수 특이적 대립단편이 확인되었다. 집 단구조에 대한 분석 결과에서 12개의 색소 및 비색소옥수수 핵심집단 계통들은 groups I, II, III, admixed group으로 구 분되었다. 본 연구결과는 옥수수 육종연구에서 색소 및 비색 소옥수수 계통들의 식별에 대한 유용한 정보를 제공할 것이다.
본 논문은 농촌진흥청 차세대바이오그린21사업(식물분자 육종사업단, PJ0080182014), 골든씨드프로젝트(식량종자사 업단, 213001-04-1-SBA10)의 지원과 강원대학교 학술연구 조성비로 연구하였음(과제번호-120140343).
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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers
Korean. J. Breed. Sci.. ;47(1):54-62.   Published online March 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.1.054
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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers
Korean. J. Breed. Sci.. ;47(1):54-62.   Published online March 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.1.054
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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers
Image Image Image Image
Fig. 1. Histogram of allele frequencies for the 1,331 alleles in 12 accessions of colored and non-colored maize lines.
Fig. 2. The Venn diagram of the number of colored and non-colored specific alleles and shared alleles.
Fig. 3. An example of a SSR profile in 12 colored and non-colored maize lines using the SSR primer, bnlg1867. ★: Specific allele to China colored maize lines, ▲: Specific allele to colored normal maize lines.
Fig. 4. UPGMA dendrogram and population structure based on the 300 SSR markers in 12 colored and non-colored maize lines. □: colored maize lines; ■: non-colored maize lines.

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Genetic Analysis of Core Sets of Colored Maize and Non-Colored Maize Inbred Lines using SSR Markers

Derivation of 12 colored and non-colored maize lines used in this study.

Code No. Entry No. Pedigree Type
1 10S4015 China collection 1-1 Colored kernel, normal corn
2 10S4026 China collection 1-4 Colored kernel, normal corn
3 10S4032 China collection 1-7 Colored kernel, normal corn
4 11CS4117 HW9 BC2 Colored kernel, waxy corn
5 11CS4124 HW9 BC2 Colored kernel, waxy corn
6 11CS7014 HW9 BC1 Colored kernel, waxy corn
7 Mo17 Mo17 Normal corn, inbred line
8 B14A B14A Normal corn, inbred line
9 HW7 HW7 Waxy corn, inbred line, black kernel
10 KW7 KW7 Waxy corn, inbred line, black kernel
11 HW9 HW9 Waxy corn, inbred line, black kernel
12 HW3 HW3 Waxy corn, inbred line, black kernel

Mean of characteristics of 300 SSR loci in 12 colored and non-colored maize lines.

Chr No. of allele MAF* GD** PIC
1 4.20 0.48 0.65 0.59
2 4.83 0.47 0.66 0.61
3 3.90 0.55 0.58 0.53
4 4.83 0.43 0.67 0.62
5 4.37 0.46 0.64 0.59
6 4.73 0.50 0.62 0.58
7 4.37 0.48 0.64 0.60
8 4.40 0.47 0.64 0.59
9 4.47 0.49 0.63 0.58
10 4.27 0.48 0.62 0.57
Total 1,331
Mean 4.44 0.48 0.64 0.59

MAF: Major Allele Frequency

GD: Genetic diversity

Mean of number of alleles and genetic diversity index for 300 SSR loci between colored and non-colored maize lines.

Chr Colored maize
 Non-colored maize
No. of Allele MAF* GD** PIC No. of Allele MAF* GD** PIC
1 3.13 0.50 0.59 0.52 3.07 0.54 0.57 0.51
2 3.00 0.55 0.54 0.49 3.30 0.50 0.60 0.54
3 2.77 0.59 0.51 0.44 2.70 0.61 0.49 0.43
4 3.23 0.52 0.57 0.51 3.67 0.46 0.64 0.58
5 2.80 0.57 0.52 0.46 3.10 0.55 0.56 0.50
6 2.53 0.64 0.44 0.39 3.67 0.48 0.63 0.58
7 2.67 0.58 0.52 0.45 3.47 0.49 0.61 0.56
8 2.53 0.60 0.48 0.41 3.67 0.47 0.64 0.58
9 2.37 0.58 0.47 0.40 3.73 0.47 0.64 0.58
10 2.73 0.57 0.51 0.45 3.30 0.52 0.59 0.53
Total 833 1,010
Mean 2.78 0.57 0.52 0.45 3.37 0.51 0.59 0.54

MAF: Major Allele Frequency

GD: Genetic diversity

Identification of colored maize specific markers among 300 SSR loci.
Table 1. Derivation of 12 colored and non-colored maize lines used in this study.
Table 2. Mean of characteristics of 300 SSR loci in 12 colored and non-colored maize lines.
Table 3. Mean of number of alleles and genetic diversity index for 300 SSR loci between colored and non-colored maize lines.
Table 4. Identification of colored maize specific markers among 300 SSR loci.

Table 1.

Table 2.

Table 3.

Table 4.

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