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

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한국, 중국 및 엘살바도르 강낭콩 (Phaseolus vulgaris L.) 유전자원의 농업 특성과 다양성 비교

Comparison of Agricultural Traits and SSR Diversity of Common Bean (Phaseouls vulgaris L. ) from Korea, China and El Salvador

Korean Journal of Breeding Science 2015;47(3):245-254.
Published online: August 31, 2015

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*Corresponding author (cym0421@korea.kr, Tel: +82-63-238-4911, Fax: +82-63-238-4909)
• Received: July 31, 2015   • Revised: September 8, 2015   • Accepted: September 22, 2015

© Korean Society of Breeding Science All rights reserved

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  • This study was conducted to compare the agricultural characteristics of total 444 common bean accessions from Korea (296), China (76), and El Salvador (72). Days to flowering were ranged from 41 to 83 days with an average of 61 days. Days to sowing to maturing were ranged from 86 to 143 days with an average of 104 days. Common beans from El Salvador tend to bloom and mature 3 to 7 days earlier than Korea and China accessions. In growth habit, over 50% of Korea and China accessions were indeterminate and climbing type, but 90.1% from El Salvador were Semi-determinate and climbing type. Qualitative traits were much different among three countries. Eighty-two percentage of immature pod colors were dark pink from El Salvador, but many of those were pale yellow from Korea (77.6%) and China (61.8%). Seed shapes were divided into four types of round, oval, cubic and kidney type, and the highest percentage of those were 30.4% with kidney type. The highest of those was 36.6% with oval type from Korea, 55.3% with kidney type from China and 79.2% with cubic type from El Salvador. Morphological characteristics of common bean from El Salvador were much different from those of Korea and China, which is necessary to collect more germplasm from its native and expand genetic diversity of common beans. Four hundred thirty-five common beans from Korea, China and El Salvador were analyzed using SSR markers. Ninety-two alleles were detected with a lowest of 6 at the BM161, BM181 and a highest of 18 at the BM154, BM160. The average polymorphism information content (PIC) was 0.72. To similar with population size among three countries, 292 Korean accessions divided four replications with 73 accessions. As a result genetic diversity was the highest of 0.73 in Chinese populations, while the lowest of 0.48 in El Salvador populations.
강낭콩(Phaseolus vulgaris L.)은 그 야생종이 미국 남서 부, 남미의 서부와 중미에 분포하고 있어 원산지를 중앙멕시 코와 과테말라로 보는 견해가 유력하며, 소립종은 중미, 대립 종은 남미에서 재배되기 시작한 것으로 알려져 있다(Phillip et al. 2011). 강낭콩은 전세계 개발도상국에서 탄수화물과 단 백질원으로 이용되는 영양학적으로 중요한 작물(Phillip et al. 2011)로, 무기질, 비타민이 풍부하고 떡소용, 밥밑콩, 협채용 등 다양한 용도로 사용되고 있다(Lee et al. 2001). 뿐만 아니 라 아시아에서는 강낭콩이 당뇨병, 심장병, 이뇨제, 류마티즘 등의 민간 의약용으로 활용되고 있어(http://www.nics.go.kr/ index.do) 최근 트렌드인 건강기능성 식품으로서도 소비자의 관심이 증가하고 있는 추세이다(Lee et al. 2001). 강낭콩의 생육적온은 20~25°C로 저온에 잘 견디고 생육기간도 비교 적 짧아 조생종은 90일 내외, 만생종은 130일 정도이므로 후 작으로 가을채소를 재배할 수 있어 작부체계상으로도 유리한 작물이다(Cheung et al. 2002, Shin et al. 2008a).
우리나라 최초의 강낭콩 품종은 일본 도입종인 Elbon을 1986년부터 생산력검정과 특성조사를 실시한 후 2년간 지역 적응시험을 거쳐 1993년 개발된 “강낭콩1호”이며, 유한신육 형, 적색종피에 신장형을 가진 풋콩용이다(Hong et al. 1993). 또한, 다양한 식생활 문화의 보급으로 인해 협채용 강낭콩의 수요가 발생하면서 황협1호가 개발되기도 하였다. 이는 풋협 을 주로 이용하므로 3월 하순에 파종하면 6~7월에 수확할 수 있는 단기성 품종으로 작부체계상 매우 유리하며 협선도 유지기간이 길어 상품성 유지에도 유리한 측면이 있다(Korea Soybean Society, 2006, Shin et al. 2008a). 신선두(경기14 호) 역시 2014년 출원된 소비자가 선호하는 선명한 자주 무 늬 종피를 가진 풋콩용 품종으로 유전자원센터에서 IT100895 를 부본으로 개발되었으며, 수확기가 7월 10일 정도로 빨라 시장성이 유리한 품종이다(GARES 2012). 강낭콩 용도는 식 생활의 변화와 함께 종실용, 협채용, 풋콩용, 가공용 등으로 다양해지고 있으나 현재 강낭콩의 재배는 대부분 재래종 또 는 도입종이 재배되고 있어 재배 안정성, 품질 및 소비자 기 호도가 높은 신품종 육성을 위하여 다양한 소재를 확보해야 할 필요성이 있다.
유전적 다양성은 유전자원의 관리와 이용에 도움을 주는 하나의 연구영역으로 분자마커를 이용하여 유전적 구조를 파 악하는 것이며, 그 중 SSR (Simple Sequence Repeat) 마커는 다른 어떤 마커보다 큰 다형성을 가지고 있어 육성품종의 유 전적 구조를 확인하기 위해 주로 이용되어 왔다(Phillip et al. 2011, Wang et al. 2006). Blair et al. (2006)은 44자원의 강 낭콩을 129개의 SSR 마커를 이용하여 안데스와 메소포타미 아 지역 자원의 genotype을 비교한 결과 안데스 자원의 genotype 다형성이 더 높다고 하였다. Gomez et al. (2005)에 따르면 농부에 의한 현지내 보존 자원과 종자은행에 의한 현지외 보 존 자원의 수량, 백립중에 대한 형태적 특성과 분자마커를 이 용한 유전적 특성을 비교한 결과 형태적 특성에서는 두 보존 방법간 유의성 있는 차이를 보이지 않았으나 유전적 다양성 분석 결과 현지외 보존자원의 다양성이 낮고 대립인자수도 적은 것으로 나타났다. 다양성의 감소는 변화하는 환경에 대 한 적응력을 약화시키며 이는 궁극적으로 생산력 감소로 이 어질 수 있다. 따라서 종자은행에서는 안전보존을 위해, 증식 시 너무 적은양의 종자를 이용해서는 안되며(Gomez et al. 2005), 형태적 특성과 더불어 유전적 특성에 대한 모니터링도 병행하여야 한다.
농업유전자원센터에 보존된 강낭콩 유전자원의 주요 수집, 도입국은 한국(1,700여점), 중국(180여점), 불가리아(170여 점), 네팔(100여점), 태국(80여점), 엘살바도르(70여점) 등이 며 총 49개국으로부터 4,200여점이 보존되어 있다. 본 실험 은 농업유전자원센터에서 보유한 중미 원산자원과 한국, 중국 등 동아시아 자원의 형태적, 유전적 특성을 비교하여 기초정 보로 이용함과 아울러 특성평가 결과의 DB 구축을 통한 정보 제공과 그에 따른 활용도 제고를 위하여 수행하였다.
유전자원의 형태적 특성
농업유전자원센터에 보존하고 있는 강낭콩 유전자원 중에서 우리나라 재래종 296자원, 중국 재래종, 육성종 등 76자원, 엘살바도르 재래종, 육성종 72자원을 포함하여 444자원을 공 시하였으며, 대조품종으로 강낭콩1호(Hong et al. 1993)와 선 두(KARES 2001)를 이용하였다. 이들 유전자원은 농업유전 자원센터에서 분양받아 이용하였으며, 엘살바도르 자원은 2013년 엘살바도르 전역에 걸쳐 수집된 것으로 수집된 전자 원을 대상으로, 한국과 중국 자원은 보존자원수를 고려하여 임의 선발하였다. 생육기간 중의 농업적 특성을 조사, 비교하 기 위하여 강낭콩 유전자원을 2014년 3월 20일 육묘폿트에 파종하여 4월 10일, 15개체씩 수원시 이목동 소재 비가림하 우스에 재식거리 90 × 50 cm, 1주 1개체로 정식하여 재배하 였다. 형태적 형질에 대한 특성조사는 식물유전자원평가기준 (RDA 1986)에 준하였다(Table 1).
Table 1.
Agricultural characteristics evaluated and their descriptors in this study.
Table 1.
Traits Methods* Rank or measurement unit Remarks
Growth habit Obs
  1. Determinate/Erect

  2. Indeterminate/Climbing

  3. Semi-determinate/Intermediate

  4. Semi-determinate/Climbing

  5. Mixed

At the end of growth
Flower color (standard) Obs
  1. White

  2. Green

  3. Purple

  4. White with purple dot or red stripe

  5. Dark purple with violet pattern

  6. Violet

  7. Mixed

Color of Petal
Days to Flowering Obs Days Days from sowing to flowering 40~50% of the plant
Days from sowing to Maturing Obs Days Days from sowing to maturing that 95% of the pod have reached final color
Pod color Obs
  1. Pale yellow

  2. Dark purple

  3. Green with purple stripe

  4. Green with red stripe

  5. Dark Pink

  6. Miscellaneous

Immatured pod color
Seed shape Obs
  • Rounded

  • Oval

  • Cubic

  • Kidney shape

  • Miscellaneous

  • Mixed

Seed shape at post-maturity
Pod length Cal Cm Pod length at post-maturity
Pod width Cal Mm Pod width at post-maturity
Seed coat color Obs
  1. White

  2. Yellow

  3. Red

  4. Violet

  5. Brown

  6. Black

  7. Pink

  8. Miscellaneous

Seed coat color at post-maturity
100 seed weight Cal Weight (g)

*Obs : Observation, Cal : Calculation.

분자생물학적 특성
분자마커를 이용한 DNA 프로파일링은 Gaitan-Solis et al. (2002)에 의해 개발된 SSR 마커 8개(Table 2)를 이용하여 한 국재래종 292, 중국 72, 엘살바도르 71자원이 포함된 435자 원에 대해 수행하였다. 파종 후 20일에 자원별 한 개체로부 터 어린잎을 채취하여 Dneasy Plant Mini Kits (Qiagen, Germantown, MD)를 이용하여 DNA를 추출하였다. 각각의 SSR 마커를 FAM, HEX, NED로 형광 표지한 후 20 ng의 강낭콩 genomic DNA, 0.2 mM의 dNTP, 50 mM의 KCl을 포함하는 1X PCR Buffer, 10 pmole의 형광표지 primer 그 리고 1unit의 Taq DNA polymerase를 혼합하여 PCR을 수행 하였다. 증폭반응은 PTC-100 thermocycler (MJ Research, Inc., Watertown, MA)를 이용하여 95°C에서 35초간 denaturation, 50~62°C에서 35초간 annealing 그리고 68°C에서 35초간 extension을 30~35회 수행하였다. 증폭산물은 ABI3130xl Genetic Analyzer (AB-PEC, Foster City, CA)를 이용하여 Cho et al. (2008)의 방법으로 분석하였다. 분석된 결과는 Genemapper 4.0 (Applied Biosystems, CA, USA.)과 powermarker3.0 (Liu & Muse 2005) 프로그램을 이용하여 유전적 다양성지 수와 heterozygosity를 산출하였다.
Table 2.
SSR markers, sequences of primers, and annealing temperatures (Tm) used for DNA profiling of common bean germplasm.
Table 2.
SSR markers Fluorescent label Forward sequences Reverse sequences Tm (°C)

BM143 HEX GGGAAATGAACAGAGGAAA ATGTTGGGAACTTTTAGTGTG 54
BM154 NED TCTTGCGACCGAGCTTCTCC CTGAATCTGAGGAACGATGACCAG 61
BM156 FAM CTTGTTCCACCTCCCATCATAGC TGCTTGCATCTCAGCCAGAATC 61
BM160 NED CGTGCTTGGCGAATAGCTTTG CGCGGTTCTGATCGTGACTTC 62
BM161 FAM TGCAAAGGGTTGAAAGTTGAGAG TTCCAATGCACCAGACATTCC 56
BM164 NED CCACCACAAGGAGAAGCAAC ACCATTCAGGCCGATACTCC 55
BM181 HEX ATGCTGCGAGTTAATGATCG TGAGGAGCAAACAGATGAGG 52
BM201 FAM TGGTGCTACAGACTTGATGG TGTCACCTCTCTCCTCCAAT 50
유전자원의 형태적 특성
숙기관련 형질 중 강낭콩 전체 자원의 평균 개화일수는 61 일, 범위는 41일에서 83일로 한국 자원의 평균 개화일수, 범 위와 같았다(Table 3). 중국 자원의 평균 개화일수는 63일, 범위는 52일에서 79일로 한국 자원의 평균 개화일수와 유사 하였으나, 변이폭은 작았다. 엘살바도르 자원의 평균 개화일 수는 58일, 범위는 54일에서 71일로 3개국 중 평균 개화일수 가 가장 짧고 변이폭도 가장 좁은 것으로 나타났다. 한국자원 의 개화일수 변이폭이 컸던 것은 자원수에 의한 측면도 있겠 으나 강낭콩이 단기성 작물로 활용되는 경우와, 자원의 일부 이겠으나, 농가 울타리에 덩굴성, 만생종으로 늦게까지 재배 하며 자가 소비하는 경우, 두 방면으로 선발되었을 가능성이 있을 것으로 보인다.
Table 3.
Days to flowering and days from sowing to maturing of common bean germplasm used this study.
Table 3.
Country Traits

Days to flowering Days from sowing to flowering

Korea Min. 41 86
Max. 83 143
Avr. 61 104

China Min. 52 90
Max. 79 118
Avr. 63 103

El Salvador Min. 54 93
Max. 71 103
Avr. 58 97

Total Avr. Standard deviation 61 103
5.95 7.21

Control cv.
Gangnangkong1 Seondu 55 96
55 96
전체 자원의 평균 생육일수는 103일, 범위는 86일에서 143 일이었으며, 한국 자원의 평균 생육일수는 104일, 범위는 86 일에서 143일로 전체 평균과 같았고 파종에서 결실까지 3개 월이 안 되는 자원부터 5개월 가량 소요되는 자원까지 변이의 폭은 57일로 매우 컸다. 강낭콩은 후작으로 가을채소 재배에 유리하기 때문에 이용하는 측면이 있음을 볼 때(Shin et al. 2008b) 생육기간 100일 미만의 자원은 조숙성 품종육종 소재 로 유용할 것으로 생각된다. 생육일수 86일인 자원은 모두 한 국 재래종으로 IT100805, IT103273, IT103353, IT103707, IT109083, IT119994, IT162766, IT194494 등 8자원이었다. 한국 자원 중 생육일수가 110일이 넘는 만생종의 비율도 10% 이상되었다(Fig. 1). 중국 자원의 평균 생육일수는 103일이었 으며, 자원의 90% 이상이 110일 이내에 성숙하였다. 엘살바 도르 자원의 평균 생육일수는 97일, 91일에서 100일 사이에 성숙하는 자원의 비율이 91.7%로 수집자원이 비교적 일제히 성숙하는 것을 알 수 있었다(Fig. 1). 생육일수의 평균과 분포 로 보아 엘살바도르 자원의 생육이 가장 빨라 조숙성 자원의 육종소재로 엘살바도르 자원이 유리할 것으로 보인다.
Fig. 1.
Frequency distribution of days to flowering and days from sowing to maturing of common bean germplasm used this study.
KJBS-47-245_F1.gif
협장과 협폭은 자원의 구분성과 더불어 자원의 용도를 구 분할 수 있는 형질 중 하나로 평균 협장은 10.9 cm, 범위는 4.2~19.5 cm로 변이가 매우 큼을 알 수 있었다(Table 4). 국 가별 평균 협장은 한국 10.1 cm, 중국 12.7 cm, 엘살바도르 12.0 cm로 중국이 가장 길고 한국이 가장 짧았다. 협장의 국 가별 분포를 보면 한국 자원은 5~10 cm에 56.2%, 10~15 cm에 42.4%로 대부분을 차지하였으며, 중국 자원은 10~15 cm 에 가장 많은 비율인 52.9%, 5~10 cm에 24.3% 였으며 15 cm 이상 자원의 비율이 22.9%로 많아 협장이 긴 자원의 비율이 다른 국가에 비해 상대적으로 높았다. 반면 엘살바도 르의 협장 분포는 협소하여 10~15 cm 범위에 93.0%의 자 원이 속하였다. 전체자원의 평균 협폭은 11.7 mm, 범위는 5.6~27.5 mm였으며, 국가별 평균 협폭은 한국 11.7 mm, 중국 12.8 mm 엘살바도르는 10.5 mm로 협폭 역시 중국자원 이 가장 컸다. 국가별 협폭의 분포는 한국 자원은 10~15 mm에 가장 많은 84.9%였고 20 mm 이상인 자원은 없었으 며, 중국은 10~15 mm 사이에 가장 많은 52.0%의 자원이 분포하였으나 20 mm 이상인 자원의 비율도 10.0%로 많았고 5~10 mm에도 24.3%의 자원이 분포하여 3개국 중 협폭의 다양성이 가장 높았다. 엘살바도르 자원은 10~15 mm (65.3%)와 5~10 mm (33.3%)에만 분포하였으며 15 mm 이 상의 협폭을 가진 자원은 없었다. 중국자원의 경우 현지 수집 시보다 국내 재배시 협장이나 협폭이 짧고 좁았다는 수집자 의 의견을 볼 때 일장, 기후, 토양조건 등에 의한 생육 차이로 생각되며, 현지에서는 협이 월등히 큰 협채용 강낭콩을 선호 하는 것으로 보인다. 협장, 협폭 조사결과를 종합하여 볼 때 한국 자원은 대체로 협이 가늘며 협당립수를 어느 정도 확보 할 수 있는 자원 위주로 선발되어 온 것을 알 수 있으며, 협장 과 협폭의 특성은 3개국의 강낭콩 용도와 선호도에 기인하는 것으로 추측되었다. 우리나라에서도 협채용 강낭콩 소비가 조 금씩 이루어지고 있으므로 협채용 신소재로 협장과 협폭이 큰 중국자원이 활용가치가 있을 것으로 생각된다.
Table 4.
Distribution of pod length and width of common bean germplasm used this study.
Table 4.
Traits Range Country Mean

Korea China El Salvador

Pod length 5 cm < 0.4% 0 0 0.2
5-10 56.2 24.3 7.0 42.9
10-15 42.4 52.9 93.0 52.4
15-20 1.0 22.9 0 4.4

Statistics (cm) Mean 10.1 12.7 12.0 10.9
Min. 4.2 7.0 9.3 4.2
Max. 18.8 19.5 14.6 19.5

Pod width 5-10 mm 11.7% 24.3 33.8 17.4
10-15 85.2 55.7 66.2 77.3
15-20 3.1 10.0 0.0 3.7
20 < 0.0 10.0 0.0 1.6

Statistics (cm) Mean 11.7 12.8 10.5 11.7
Min. 8.1 6.7 5.6 5.6
Max. 15.9 27.5 13.4 27.5
두과작물에서 100립중은 그 편차로 인해 품종을 구분하는 중요한 특성이 되기도 하는데 강낭콩 역시 100립중 편차가 큰 작물 중 하나이다. 평균 100립중은 40.9 g, 범위는 18.1~ 69.3 g이었으며 국가별로 한국 자원은 평균 44.3 g, 18.1~ 69.3 g, 중국 자원은 평균 42.7 g, 19.2~67.6 g, 엘살바도르 자원은 평균 26.3 g, 18.5~32.1 g으로 한국과 중국 자원의 평균 100립중과 범위는 유사하였으나 엘살바도르 자원의 평 균 100립중은 한국, 중국보다 작고 그 범위도 비교적 좁았다. 본 결과는 100립중 범위가 19~71 g 사이라는 Choi et al. (2012)의 결과와 유사하였으며, 강낭콩은 품종이 많고 종피색 이 다양하며 100립중이 20~60 g 내외라고 한 Lee et al. (2001)의 보고보다 100립중의 범위가 큰 것으로 조사되어 꾸 준한 유전자원의 수집, 도입이 유전적 다양성 증대에 기여한 것으로 생각된다. 100립중은 46~60 g의 범위에 자원의 41.1%가 포함되어 가장 많이 분포하였으며, 31~45 g의 범 위에 34.3%, 30 g 미만에 22.9%로 비교적 고르게 분포하는 것으로 나타났다. 다만, 60 g 이상의 대립종은 1.7%에 불과 하였다. 국가별로 한국과 중국 자원은 46~60 g사이의 대립 종 자원이 각각 50.9%, 44.6%로 가장 많고 엘살바도르 자원 은 30 g 미만의 소립자원이 90% 이상으로 많아 중미지역은 소립종 위주로 발달했다는 보고(Phillip et al. 2011)와 일치 하는 것으로 나타났다.
질적형질 중 생육습성은 그 작물의 재배에 있어서 파종기 와 재배적기를 선택하는데 중요한 형질로 신육형이 유한형이 며 직립형 자원이 35.7%, 무한형이며 덩굴성 자원이 46.4% 로 가장 많았고, 신육형이 중간형이고 초형 역시 중간형인 자 원은 0.5%, 중간형이며 덩굴성인 자원은 16.1%를 차지하였 다. 생육습성 역시 국가별로 차이를 보여, 한국 자원은 유한형과 무한형의 비율이 유사하였으나, 중국 자원은 무한형이 60.5% 로 많았고, 엘살바도르 자원은 대부분 중간형으로 90.1%였 다. 수집자에 따르면 엘살바도르 수집 자원은 현지에서는 키 가 작은 유한 직립형이라고 하였으나 국내 재배시 중간형 특 성을 보여 일장 및 기후조건에 의한 생육차로 보였다. 우리나 라의 강낭콩은 가을채소를 심기 전에 단작으로 재배하는 경 우가 많아 생육기가 짧은 직립형을 선호하는 경향이 있으나 수량이 적다는 단점이 있다. 또한, 무한형은 만생종으로 작기 가 길고 우거져서 관리가 어려운 단점이 있으므로 중간형 자 원이 개발될 경우 유한형과 무한형의 단점을 보완할 수 있는 좋은 육종소재가 될 것으로 보인다. 실제로 엘살바도르 수집 자원의 생육기간은 평균 97일로 한국, 중국 자원보다 빠르고 대조품종으로 사용된 강낭콩1호와 선두보다 하루 늦어 육종 소재로서 유용할 것으로 생각된다.
강낭콩의 화색은 기판과 익판으로 구분되며, 기판의 화색 은 보라색이 36.4%로 가장 많고, 보라색 줄무늬가 있는 백색 (28.1%), 자주색(13.1%), 백색(11.3%)순이었다. 국가별로 한 국 자원은 보라색 44.6%, 중국 자원은 보라색 38.2%, 자주색 32.9%, 엘살바도르 자원은 보라색 줄무늬가 있는 백색(86.1%) 에 집중되었다. 화색의 익판은 보라색이 59.3%로 가장 많았 으며 국가별로 한국(76.5%)과 중국 자원(42.1%)은 보라색이 많고, 엘살바도르 자원은 백색의 비율이 91.7%로 대부분이었 다. 강낭콩 협색은 미성숙 협색과 성숙협색으로 나누어지며, 본 연구결과에서 미성숙 협색은 크게 연노랑, 녹색바탕에 적 색 줄무늬, 진분홍색으로 구분이 되었으며, 전체적으로 연노 랑색 비율이 63.3%로 가장 많았고, 국가별로 한국과 중국 자 원의 협색은 연노랑이 주였으나(각각 77.6%, 61.8%), 엘살바 도르 자원은 81.9%가 진홍색으로 다른 자원들과 확연히 구분 되었다. 엘살바도르 자원의 미성숙 협색은 추후 안토시아닌 분석을 통하여 천연염료로서 활용가능성을 검토할 필요가 있 을 것으로 보인다. 또한 미성숙 협색이 진홍색으로 화려하고, 신육형이 덩굴성인 특성이 있어 울타리 경관용으로도 가치가 있을 것으로 보인다. 성숙협색은 노랑, 갈색, 흑녹색, 흑갈색 으로 구분이 되었으며, 흑녹색이 67.1%로 가장 많았고 국가 별로도 한국, 중국, 엘살바도르 자원 모두 흑녹색의 비율이 각 각 53.9%, 63.6%, 79.2%로 가장 많아 성숙한 협색에서는 큰 차이를 볼 수 없었다.Table .5,6
Table 5.
Distribution of 100 seed weight of common bean germplasm used this study. unit : %
Table 5.
Country 100 seed weight Statistics

30 g > 31-45 g 46-60 g 61 g < Min. (g) Max. (g) Mean (g)

Korea 6.7 40.8 50.9 1.5 18.1 69.3 44.3
China 17.6 33.8 44.6 4.1 19.2 67.6 42.7
El Salvador 90.0 10.0 0 0 18.5 32.1 26.3

Total 22.9 34.3 41.1 1.7 18.1 67.6 40.9
Table 6.
The distribution of Growth habit of common bean germplasm used this study. unit : %
Table 6.
Growth habit KOR CHN SLV Total

Determinate/Erect 46.4 27.6 0.0 35.7
Indeterminate/Climbing 52.6 60.5 5.6 46.4
Semi-determinate/Intermidiate 0.0 0.0 2.8 0.5
Semi-determinate/Climbing 0.7 6.6 90.1 16.1
Mixed 0.3 5.3 1.5 1.3

*Growth habit of check variety : Gangnangkong 1 (Determinate/Erect), Seondu (Determinate/Erect).

종피색은 전체적으로 적색자원이 44.5%로 가장 많았고 갈 색자원 22.7%, 백색자원 10.2%, 검정색 5.7% 순이었다. 국 가별로 한국과 엘살바도르 수집자원은 적색자원의 비율이 42.1%와 83.3%로 가장 많았고, 중국은 갈색자원이 36.8%로 많았다. 엘살바도르에서 강낭콩은 고물용으로 많이 활용된다 고 하며 이는 적색종피의 자원이 많은 이유일 것으로 보인다. 종자모양은 둥근형, 계란형, 입방형, 신장형으로 나뉘어졌으 며, 신장형이 30.4%로 가장 많고 계란형(27.4%)과 입방형 (21.0%) 순이었다. 한국은 계란형(36.6%), 신장형(29.7%)순, 중국은 신장형(55.3%), 엘살바도르는 입방형(79.2%)이 많아 종자모양 역시 국가별 차이를 보였다. 질적형질인 생육습성, 미성숙 협색, 화색, 종자모양 등에서 엘살바도르 자원과 동아 시아 자원의 분포는 상당한 차이가 있었다. 엘살바도르는 강 낭콩 원산지임에도 불구하고 한국, 중국의 자원들과 달리 특 성이 어느 한 형질에 집중되는 경향이 있어 엘살바도르에서 강낭콩의 용도, 선호도와 같은 추가 정보를 통한 분석이 필요 할 것으로 보인다. 그러나, 생육습성과 종자모양, 미성숙 협색 등에서 동아시아에서는 거의 볼 수 없는 형질을 가지고 있어 강낭콩 원산 자원을 추가 확보하여 다양성을 높일 필요가 있다.Fig .2
Fig. 2.
Frequency distribution of qualitative traits including colors of flower, seed coat color, seed shape and pod color in common bean germplasm used this study.
KJBS-47-245_F2.gif
분자생물학적 특성
각국 자원의 유전적 다양성을 비교하기 위하여 한국 292자 원, 중국 72자원, 엘살바도르 71자원을 포함한 435자원에 대 해 8개의 SSR 마커를 이용하여 DNA 프로파일링을 수행하 였으며, 그 결과 총 92개의 allele가 확인되었고 마커에 따라 allele 수는 최소 6개(BM161, BM181)에서 18개(BM160, BM154), 평균 11.5개로 나타났다(Table 7). Gomez et al. (2005)에 따르면 8개 강낭콩 자원을 대상으로 7개의 마커를 이용하여 분석한 결과 유전자좌당 allele 수는 2개에서 9개, 평균 4.3개를 가지는 것으로 조사되었으며, Phillip et al. (2011)은 181점의 멕시코 및 중남미 지역에서 수집된 강낭콩 재래종 자원을 58개의 SSR 마커를 이용하여 분석한 결과 유 전자좌당 allele의 수는 2개에서 8개 사이라고 보고한 바 있는 데 본 연구결과에서는 이보다 많은 allele가 확인되었다.
Table 7.
SSR markers used in this study, their associated linkage groups, major allele frequency, allele size range, the number of alleles per locus, gene diversity, heterozygosity, and polymorphism information content (PIC).
Table 7.
Marker Linkage group* Major allele frequency Allele size range Allele no. per locus Gene diversity Heterozygosity PIC

BM143 2 0.24 111-156 14 0.85 0.024 0.83
BM154 9 0.37 208-270 18 0.79 0.026 0.77
BM156 2 0.19 214-277 16 0.89 0.007 0.88
BM160 7 0.26 181-257 18 0.87 0.009 0.85
BM161 4 0.56 177-189 6 0.57 0.000 0.50
BM164 2 0.41 135-175 7 0.68 0.002 0.62
BM181 3 0.31 182-204 6 0.78 0.023 0.75
BM201 7 0.64 94-112 7 0.55 0.005 0.52

Mean 0.37 11.5 0.75 0.014 0.72

*The assignment of SSR markers to linkage groups is based on Gaitan-Solis et al. (2002).

마커별 PIC는 BM156 마커가 0.88로 가장 높았고, BM161 마커가 0.50로 가장 낮게 나타났으며 평균 0.72로 비교적 높 았다. 이는 불가리아 수집자원과 한국재래종 강낭콩 자원을 대상으로 다양성 분석한 PIC값 0.62 (Choi et al. 2012) 보다 높았는데, 원산지인 엘살바도르 자원이 추가되어 다양성을 높인 결과일 것으로 생각된다. 분석에 이용된 8개 마커의 heterozygosity는 0.0~2.6%로 평균 1.4%를 나타냈다.
각 수집 국가별 다양성 분석 결과를 비교하기 위하여 중국, 엘살바도르는 72자원, 71자원을 공시하였으며, 한국자원은 다른 집단과 크기를 유사하게 하기 위해 73자원씩 4반복하여 그 평균을 이용하였다. 그 결과 한국자원의 평균 allele 수는 7.5개, 중국 8.3개, 엘살바도르 7.0개로 중국자원의 allele 수 가 많은 것으로 나타났다. 중국자원의 allele 수가 많았던 것 은 중국이 대륙으로 자원의 분포범위가 폭 넓다는 것과 다양 한 식습관으로 인해 다각적으로 활용되어왔기 때문일 것으로 보인다. 유전적 다양성 지수와 PIC도 중국 자원이 각각 0.73 과 0.69로 가장 높았으며, 엘살바도르 자원이 0.48, 0.45로 가 장 낮음을 알 수 있었다(Table 8). Lee et al. (2009)이 불가 리아 남서부지역 수집 재래종과 한국 재래종 강낭콩의 유전 적 다양성 분석 결과 PIC가 0.5639이었던 결과나 Blair et al. (2002)이 강낭콩 원산지인 중남미 자원 42점을 대상으로 분 석한 결과인 0.594보다 한국과 중국자원의 PIC는 높았으나 엘살바도르 자원에서는 매우 낮은 것으로 나타났다. 일반적으 로 유전적 다양성은 원산지 지역에서 수집된 유전자원에서 더 높다고 알려져 있는데 본 연구에 사용된 엘살바도르 수집 자원의 다양성 지수와 PIC 값이 낮았던 것은 고물용과 같이 한 방향으로 선발되어 진화되었을 가능성이 있다. 또한, 성숙 일수, 화색, 협색, 백립중, 종피색, 신육형 등 농업특성에서도 특정한 형질에 집중되어 있었는데, 이는 엘살바도르가 원산지 이나 우리나라 국토의 1/5 정도로 좁고 원산지의 극히 일부이 기 때문일 수 있을 것으로 보인다. 그러나 중미 원산자원이 동아시아 자원이 가진 농업특성과는 매우 다른 양상을 보이 고 있어 농업유전자원센터의 다양성 확보를 위하여 강낭콩 원산자원을 확보해야 할 필요성은 충분하며, 추후 보다 많은 원산자원과 마커를 선발하여 심도 있는 유전적 다양성 분석 을 수행해야 할 것으로 보인다.
Table 8.
Major allele frequency, the number of alleles, gene diversity, heterozygosity, polymorphism information content (PIC) based on the origin of common bean germplasms.
Table 8.
Origin Major Allele Frquency Allele no. per locus Gene Diversity Heterozygosity PIC

Korea 0.44 7.5 0.68 0.015 0.64
China 0.39 8.3 0.73 0.015 0.69
El Salvador 0.65 7.0 0.48 0.013 0.45

Mean 0.49 7.6 0.63 0.014 0.59
본 연구는 한국, 중국, 엘살바도르 강낭콩 유전자원을 대상 으로 농업특성을 조사하고, 분자마커를 이용하여 유전적 다양 성을 분석하였으며 결과는 다음과 같다. 첫째, 개화일수 전체 평균은 61일, 범위는 41일에서 83일, 생육일수는 평균 104일, 범위는 86일에서 143일 사이였다. 엘살바도르 자원의 평균 생육일수는 97일로 한국, 중국의 104일과 103일보다 6~7일 빨랐다. 둘째, 협장은 전체 평균 10.9 cm, 범위는 4.2~19.5 cm, 중국 자원의 평균 협장은 12.7 cm로 가장 길고 범위도 7.0~19.5 cm로 넓었다. 협폭의 전체 평균은 11.7 mm, 범위 는 5.6~27.5 mm, 중국 자원의 평균 협폭은 12.8 mm, 범위는 6.7~27.5 mm로 3개국 중 가장 넓었다. 셋째, 100립중의 전 체 평균은 40.9 g, 범위는 18.1~69.3 g, 한국과 중국 자원의 평균 100립중은 각각 44.3 g, 42.7 g으로 유사하였으나 엘살 바도르 자원의 평균 100립중은 26.3 g으로 매우 작았으며 분 포범위도 18.5~32.1 g으로 좁았다. 넷째, 신육형은 유한형이 며 직립형 자원이 35.7%, 무한형이며 덩굴성인 자원이 46.4%였으며, 신육형이 중간형이고 덩굴성 자원이 16.1%로 나타났다. 한국 자원은 유한형과 무한형의 비율이 유사하였으 며, 중국 자원의 신육형은 무한형이며, 덩굴성 자원이 60.5%, 엘살바도르는 중간형이며, 덩굴성 자원이 90.1%로 대부분을 차지하였다. 다섯째, 종피색은 전체적으로 적색자원이 평균 44.5%로 가장 많았으며, 특히 엘살바도르는 적색자원이 83.3%로 대부분이었고, 종자모양은 신장형이 평균 30.4%로 가장 많았으나, 한국자원은 계란형(36.6%), 중국자원은 신장 형(55.3%), 엘살바도르는 입방형(79.2%)이 많아 국가별 차이 를 보였다. 여섯째, 한국 자원 292점, 중국 자원 72점, 엘살바 도르 자원 71점 등 435점을 대상으로 8개의 SSR 마커를 이 용한 유전적 다양성 분석을 수행하였으며, 그 결과 총 92개의 allele가 확인되었고, gene diversity 와 PIC로 볼 때 중국 자 원이 각각 0.73, 0.69로 가장 높았으며, 엘살바도르 자원은 0.48과 0.45로 가장 낮은 것으로 나타났다.
본 연구는 농촌진흥청 농업과학기술연구 개발사업(과제번 호: PJ01010603)의 지원에 의해 이루어졌습니다.
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Comparison of Agricultural Traits and SSR Diversity of Common Bean (Phaseouls vulgaris L. ) from Korea, China and El Salvador
Korean. J. Breed. Sci.. 2015;47(3):245-254.   Published online September 30, 2015
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Comparison of Agricultural Traits and SSR Diversity of Common Bean (Phaseouls vulgaris L. ) from Korea, China and El Salvador
Korean. J. Breed. Sci.. 2015;47(3):245-254.   Published online September 30, 2015
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Comparison of Agricultural Traits and SSR Diversity of Common Bean (Phaseouls vulgaris L. ) from Korea, China and El Salvador
Image Image
Fig. 1. Frequency distribution of days to flowering and days from sowing to maturing of common bean germplasm used this study.
Fig. 2. Frequency distribution of qualitative traits including colors of flower, seed coat color, seed shape and pod color in common bean germplasm used this study.
Comparison of Agricultural Traits and SSR Diversity of Common Bean (Phaseouls vulgaris L. ) from Korea, China and El Salvador

Agricultural characteristics evaluated and their descriptors in this study.

Traits Methods* Rank or measurement unit Remarks
Growth habit Obs

Determinate/Erect

Indeterminate/Climbing

Semi-determinate/Intermediate

Semi-determinate/Climbing

Mixed

At the end of growth
Flower color (standard) Obs

White

Green

Purple

White with purple dot or red stripe

Dark purple with violet pattern

Violet

Mixed

Color of Petal
Days to Flowering Obs Days Days from sowing to flowering 40~50% of the plant
Days from sowing to Maturing Obs Days Days from sowing to maturing that 95% of the pod have reached final color
Pod color Obs

Pale yellow

Dark purple

Green with purple stripe

Green with red stripe

Dark Pink

Miscellaneous

Immatured pod color
Seed shape Obs

Rounded

Oval

Cubic

Kidney shape

Miscellaneous

Mixed

Seed shape at post-maturity
Pod length Cal Cm Pod length at post-maturity
Pod width Cal Mm Pod width at post-maturity
Seed coat color Obs

White

Yellow

Red

Violet

Brown

Black

Pink

Miscellaneous

Seed coat color at post-maturity
100 seed weight Cal Weight (g)

*Obs : Observation, Cal : Calculation.

SSR markers, sequences of primers, and annealing temperatures (Tm) used for DNA profiling of common bean germplasm.

SSR markers Fluorescent label Forward sequences Reverse sequences Tm (°C)

BM143 HEX GGGAAATGAACAGAGGAAA ATGTTGGGAACTTTTAGTGTG 54
BM154 NED TCTTGCGACCGAGCTTCTCC CTGAATCTGAGGAACGATGACCAG 61
BM156 FAM CTTGTTCCACCTCCCATCATAGC TGCTTGCATCTCAGCCAGAATC 61
BM160 NED CGTGCTTGGCGAATAGCTTTG CGCGGTTCTGATCGTGACTTC 62
BM161 FAM TGCAAAGGGTTGAAAGTTGAGAG TTCCAATGCACCAGACATTCC 56
BM164 NED CCACCACAAGGAGAAGCAAC ACCATTCAGGCCGATACTCC 55
BM181 HEX ATGCTGCGAGTTAATGATCG TGAGGAGCAAACAGATGAGG 52
BM201 FAM TGGTGCTACAGACTTGATGG TGTCACCTCTCTCCTCCAAT 50

Days to flowering and days from sowing to maturing of common bean germplasm used this study.

Country Traits

Days to flowering Days from sowing to flowering

Korea Min. 41 86
Max. 83 143
Avr. 61 104

China Min. 52 90
Max. 79 118
Avr. 63 103

El Salvador Min. 54 93
Max. 71 103
Avr. 58 97

Total Avr. Standard deviation 61 103
5.95 7.21

Control cv.
Gangnangkong1 Seondu 55 96
55 96

Distribution of pod length and width of common bean germplasm used this study.

Traits Range Country Mean

Korea China El Salvador

Pod length 5 cm < 0.4% 0 0 0.2
5-10 56.2 24.3 7.0 42.9
10-15 42.4 52.9 93.0 52.4
15-20 1.0 22.9 0 4.4

Statistics (cm) Mean 10.1 12.7 12.0 10.9
Min. 4.2 7.0 9.3 4.2
Max. 18.8 19.5 14.6 19.5

Pod width 5-10 mm 11.7% 24.3 33.8 17.4
10-15 85.2 55.7 66.2 77.3
15-20 3.1 10.0 0.0 3.7
20 < 0.0 10.0 0.0 1.6

Statistics (cm) Mean 11.7 12.8 10.5 11.7
Min. 8.1 6.7 5.6 5.6
Max. 15.9 27.5 13.4 27.5

Distribution of 100 seed weight of common bean germplasm used this study. unit : %

Country 100 seed weight Statistics

30 g > 31-45 g 46-60 g 61 g < Min. (g) Max. (g) Mean (g)

Korea 6.7 40.8 50.9 1.5 18.1 69.3 44.3
China 17.6 33.8 44.6 4.1 19.2 67.6 42.7
El Salvador 90.0 10.0 0 0 18.5 32.1 26.3

Total 22.9 34.3 41.1 1.7 18.1 67.6 40.9

The distribution of Growth habit of common bean germplasm used this study. unit : %

Growth habit KOR CHN SLV Total

Determinate/Erect 46.4 27.6 0.0 35.7
Indeterminate/Climbing 52.6 60.5 5.6 46.4
Semi-determinate/Intermidiate 0.0 0.0 2.8 0.5
Semi-determinate/Climbing 0.7 6.6 90.1 16.1
Mixed 0.3 5.3 1.5 1.3

*Growth habit of check variety : Gangnangkong 1 (Determinate/Erect), Seondu (Determinate/Erect).

SSR markers used in this study, their associated linkage groups, major allele frequency, allele size range, the number of alleles per locus, gene diversity, heterozygosity, and polymorphism information content (PIC).

Marker Linkage group* Major allele frequency Allele size range Allele no. per locus Gene diversity Heterozygosity PIC

BM143 2 0.24 111-156 14 0.85 0.024 0.83
BM154 9 0.37 208-270 18 0.79 0.026 0.77
BM156 2 0.19 214-277 16 0.89 0.007 0.88
BM160 7 0.26 181-257 18 0.87 0.009 0.85
BM161 4 0.56 177-189 6 0.57 0.000 0.50
BM164 2 0.41 135-175 7 0.68 0.002 0.62
BM181 3 0.31 182-204 6 0.78 0.023 0.75
BM201 7 0.64 94-112 7 0.55 0.005 0.52

Mean 0.37 11.5 0.75 0.014 0.72

*The assignment of SSR markers to linkage groups is based on Gaitan-Solis et al. (2002).

Major allele frequency, the number of alleles, gene diversity, heterozygosity, polymorphism information content (PIC) based on the origin of common bean germplasms.

Origin Major Allele Frquency Allele no. per locus Gene Diversity Heterozygosity PIC

Korea 0.44 7.5 0.68 0.015 0.64
China 0.39 8.3 0.73 0.015 0.69
El Salvador 0.65 7.0 0.48 0.013 0.45

Mean 0.49 7.6 0.63 0.014 0.59
Table 1. Agricultural characteristics evaluated and their descriptors in this study.

Obs : Observation, Cal : Calculation.

Table 2. SSR markers, sequences of primers, and annealing temperatures (Tm) used for DNA profiling of common bean germplasm.
Table 3. Days to flowering and days from sowing to maturing of common bean germplasm used this study.
Table 4. Distribution of pod length and width of common bean germplasm used this study.
Table 5. Distribution of 100 seed weight of common bean germplasm used this study. unit : %
Table 6. The distribution of Growth habit of common bean germplasm used this study. unit : %

Growth habit of check variety : Gangnangkong 1 (Determinate/Erect), Seondu (Determinate/Erect).

Table 7. SSR markers used in this study, their associated linkage groups, major allele frequency, allele size range, the number of alleles per locus, gene diversity, heterozygosity, and polymorphism information content (PIC).

The assignment of SSR markers to linkage groups is based on Gaitan-Solis et al. (2002).

Table 8. Major allele frequency, the number of alleles, gene diversity, heterozygosity, polymorphism information content (PIC) based on the origin of common bean germplasms.