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국내 육성 벼 243 품종의 형태적 특성에 따른 품종 분류

Classification of Korean Rice Varieties Based on Agro-morphological Traits

Me-Sun Kim1, Hye-Jung Lee1,2, Dal-A Yu1, Jae-Young Song1, Marjohn Nino1, Franz Nogoy1, Joonki Kim1, Yoon-Sup So1, Yong-Gu Cho1,*
The Korean Journal of Breeding Science 2016;48(3):254-270.
Published online: August 31, 2016

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*Corresponding author (ygcho@cbnu.ac.kr, +82-43-261-2514, +82-43-273-2242)
• Received: July 8, 2016   • Accepted: August 8, 2016

© The Korean Society of Breeding Science

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  • The International Union for the Protection of New Varieties of Plants (UPOV) promotes an effective system of plant variety protection and encourages the development of new varieties of plants. This international convention was initiated to standardize the system efforts and strengthen policy. The establishment of cultivar discrimination system is very important to distinguish varieties between domestic and foreign agricultural products. It is necessary for the protection of breeders’rights. In addition, it will help for more efficient and quality management of plant breeding. This study was conducted to identify and group rice varieties based on agro-morphological characteristics such as plant height, panicle length, number of tillers, culm length, leaf length, leaf width, leaf pigments and flag leaf angles. Using these parameters, statistical analysis classified a total of 243 rice varieties bred in Korea into four groups. Most rice varieties did not exhibit anthocyanin pigments on the leaves particularly on the first leaf, leaf blade, leaf sheath and auricle, except for varieties classified as black rice. Results of phylogenetic and principal component analysis (PCA) indicated that these varieties formed three largely distinct clusters according to their ecotype and morphological differentiation. This result would be useful in rice varietal identification for the protection of breeders’variety rights.
우리나라에서 벼 품종의 육종은 1950년대에 교배육종을 통하 여 다수성 품종 육성에 주력하면서 내도복성과 내병성 증진에 노력해왔고, 1970년대에는 japonica와 indica 벼의 원연교잡으 로 다수성과 내도복성 및 내병충성 증대에 모든 육종적 노력을 집중하는 한편, 통일형 품종의 결점인 내냉성, 내탈립성, 그리고, 미질개선을 위해 노력해왔다. 1980년대의 큰 냉해 피해 이후에 는 내냉성 강화에 힘을 기울였고, 쌀의 과잉생산과 소비자들의 양질미 선호도가 증가함에 따라 쌀의 미질개선에도 힘을 기울였 다(Lee et al. 2012). 그 후에는 japonica 벼 품종의 초형 개선에 의한 수량성과 안정성 증대에 주력하였다. 그리고, 쌀의 소비증 국내 육성 벼 243 품종의 형태적 특성에 따른 품종 분류대 및 부가가치 증진을 위한 가공적성 증진, 병충해 및 기상재해 복합저 항성의 유전적 보완이 강화되었다. 1990년대 이후에는 쌀의 품질이 우수하고 밥맛이 좋은 양질미의 수량성과 안정성 증대를 위해 노력함과 동시에 가공용 벼 품종개발 등 품종의 다양화에 주력하여 왔다(Kang et al. 2004, Breseghello 2013). 최근에는 WTO와 FTA에 의하여 쌀 시장의 개방에 대비하여 국제경쟁력 강화를 위한 고품질 품종육성에 주력하면서 남북통일에 대비하 고 식량안보의 확보를 위한 쌀 수량성 증대연구와 병해충과 재해에 복합저항성을 지닌 품종개발을 지속하고 있다(Kim 2013).
벼 육종가가 새로운 품종을 육성하기 위해서는 10~15년의 기간이 소요되는 등 많은 시간과 노력이 투여되는 반면, 일단 새로운 품종이 육성되어 공개되면 많은 경우 다른 사람들에 의해 쉽게 재생산되어 그것을 개발한 육종가에게 투자에 대한 적절한 보상의 기회를 박탈해 버림으로써 의욕을 상실하게 만든 다(Choi et al. 2004). 따라서 식물육종 분야에 있어서 개인 육종가, 발명자에게 그 신품종의 개발에 이르기까지의 시간과 노력, 재정적인 면 등에 있어서 그 대가로서 지식재산권을 부여하 여 보호할 필요가 있는 것이다(Park et al. 2009). 이 밖에도 종자에 대해 지식재산권을 부여함으로써 정부 또는 공공부분의 역할을 줄이고 민간부문을 통해 품종육성 및 종자생산을 할 수 있다는 점 등을 지적하기도 한다.
1998년부터 도입된 품종보호제도는 식물 신품종 육성자의 권리를 법적으로 보장하여 주는 지적재산권의 한 형태로 육성자 에게 배타적인 상업적 독점권을 부여하여 품종 개발을 촉진하기 위한 제도이다(Seong et al. 2009).
품종보호제도의 규범화는 신품종의 육성을 촉진하고 우수품 종의 이용을 확대하여 농업의 생산성을 증대시킬 수 있는 긍정적 인 효과가 기대되며, 이와 같이 법적으로 식물품종에 대한 보호를 의무화하려는 추세가 전세계적으로 확대되고 있다(Choi 2004, Garris et al. 2005).
최근 선진국들은 신품종 종자산업을 국가경쟁력의 새로운 원천으로 인식하여 지원을 강화하고 있다(Osteen et al. 2012, SCBD 2005). 그러나 우리나라는 ‘종자산업법’에서 신품종 육성 자의 권리를 법적으로 보장하기 위해 식물 신품종보호제도를 채택하고 있으나 품종보호에 관한 국내에서의 논의는 활발하지 않다. 또한, 식물에 대한 국내 유통품종 중 법정종자를 제외한 품종의 특성 데이터베이스를 구축하여 관리하는 노력이 부족하 며, 품종 간 유전적 최소거리(Minimum Distance)에 대한 법정 분쟁 및 환경과 외형의 차이에 따른 특성별 차이에 대한 연구가 미흡하다. 벼에서는 1984년부터 1994년까지 수많은 형태적 돌 연변이 특성들의 연관군지도를 작성하여 그들의 염색체별 위치 를 보고한 바 있는데(Kinoshita 1984~1994, Causse et. al. 1994), 이러한 형태적 특성의 변이를 이용하면 수많은 유사한 벼의 품종을 구분하는 데에 매우 효과적으로 이용될 수 있을 것으로 생각된다.
우리나라 국가기관에서는 18 개 작물에 대한 DNA Profile의 데이터베이스를 구축하여 다방면에 활용하고 있으나, 국제신품 종보호동맹 (International Union for the Protection of New Varieties of Plants, UPOV)의 사례와 같이 국가적이고 표준화된 데이터베이스의 구축은 미흡한 실정으로 차후에 분석결과의 신 뢰도 및 공정성에 대한 문제점이 발생할 수 있을 것으로 예상된다.
따라서, 본 연구에서는 우리나라에서 품종으로 육성되어 유통 되고 있는 벼 품종의 외형의 차이에 따른 품종별 형태적 특성을 농촌진흥청 및 국립종자원의 조사기준에 따라서 조사하였고 (RDA 2003a) 벼 품종 간 유전적 거리를 탐색하여 각 품종의 형질 측정을 통해 데이터베이스를 구축하여 표준품종을 선정하 고 품종의 형태적 특성뿐만 아니라 DNA profiling에 의하여 품종에 대한 구분체계를 확립할 수 있는 기초 정보를 제공하고자 하였다.
본 보고에서는 국내에서 육성하여 국립종자원을 통하여 보급 되고 있는 243개 벼 품종에 대하여 특히 조사가 가능한 벼의 형태적 특성을 질적형질과 양적형질 모두를 세밀하게 조사하여 주성분 분석 및 군집 분석을 수행하였고 그를 통하여 품종군을 분류하였으며, 초형, 이삭 및 현미의 모양을 사진을 찍어 직접 비교에 활용할 수 있게 제공하였다.
시험재료 및 재배법
시험재료는 국내에서 육성된 벼 품종 중 1982년부터 2013년 까지 국립종자원을 통하여 보급된 243개 벼 품종이 사용되었다 (Table 1). 벼 품종은 2013년국내 육성 벼 243 품종의 형태적 특성에 따른 품종 분류과 2014년에 충북대학교 시험포장에 이앙 하여 관찰, 조사하였다. 품종 당 1주 1본씩, 16주/열, 3열로 이앙 하였으며 물 관리와 비료의 시용 및 병, 해충의 방제는 농촌진흥 청 벼 표준재배법에 준하여 수행하였다(RDA 2003a).
Table 1.
List of 243 rice varieties bred in Korea from 1982 to 2013.
Table 1.
조사항목
벼의 형태적 특성 조사를 위하여 양 끝에 심겨진 3개체씩을 배제하고 안쪽에 심겨진 10주를 기준으로 하여 3반복으로 농 촌진흥청 조사기준 및 국립종자원에서 규정한 벼 표준조사방 법에 따라 조사하였다(RDA 2003a). 형태적 특성 조사를 크게 잎, 줄기, 출수기로 나누어 조사하였으며, 질적형질은 육안조사 를 실시하였고, 양적형질은 1000 mm 금속 자를 이용하여 측량 조사를 실시하였다(Table 2). 조사항목은 초장, 간장, 제3엽기 에 제1엽의 안토시아닌 색소 유무, 잎의 녹색정도 및 안토시아 닌색소 유무, 잎혀의 모양 및 색, 잎몸의 길이 및 너비를 조사하 였으며, 잎몸의 길이 및 너비는 출수 직후에 상위 3번째 엽의 길이를 측정하였다. 줄기는 지엽의 자세 및 굵기, 길이, 모양을 조사하였으며, 이때 지엽의 자세는 지엽의 만곡성 정도를 가지 고 조사하였고 줄기의 굵기는 주간 엽신의 폭이 가장 넓은 부위 를 조사하였다. 출수기는 조사개체의 50%가 출수하였을 때를 출수기로 하였다.
Table 2.
Agronomic traits investigated in the morphological analysis of rice varieties.
Table 2.
Characteristics Subsection Stage
Qualitative trait
1st leaf Color Nursery stage
Leaf Intensity of green color Maximum tiller number stage
Distribution of anthocyanin
Ligule Color, Shape Maximum tiller number stage
Flag leaf Attitude Heading stage, Yellow ripe stage
Culm Thickness, Attitude Dough ripe stage, Maturity stage
Quantitative trait
Leaf blade Length, Width (cm) Maximum tiller number stage
Culm Length (cm) Dough ripe stage, Maturity stage
Time of heading Time of heading (Day) 50% heading of rice
Panicles per plant (No.)
Panicle Length (cm) Dough ripe stage, Maturity stage
통계분석
통계프로그램 SAS version 9.2 (SAS Institute Inc.)과 Multibase program (http://www.numericaldynamics.com)을 이용하였으며 2013년도와 2014년도에 243개 품종으로부터 얻 어진 데이터를 분산분석을 통해 유의성을 검정하였으며, 벼 243 개 품종들간의 형태적 특성에 기초한 품종군을 분류하고자 시각 적으로 가장 두드러지는 형태적 특성인 출수기를 비롯하여 초장, 간장, 수장, 분얼수 및 잎몸의 너비와 길이 등 총 7개의 형질을 이용하여 주성분 분석과 군집분석을 수행하였다.
형태적 특성
(1)

(1) 잎의 길이와 너비

벼 243 품종의 잎 형태적 특성을 비교 분석하기 위하여 잎몸의 길이 및 너비를 조사하였다.
잎몸의 길이 및 너비는 출수 직후에 상위 3번째 엽의 길이를 측정하였으며 개화시기별로 나타낸 결과는 Table 3과 같다. 잎몸 길이에서 조생종은 월백이 23.44 cm 국내 육성 벼 243 품종의 형태적 특성에 따른 품종 분류로 가장 짧았고 중모1012가 47.88 cm로 가장 길었으며 평균은 30.96 cm를 나타내었다. 중생종은 큰섬이 21.76 cm로 가장 짧았고 소비가 59.10 cm로 가장 길었으 며 평균은 39.74 cm로 나타났으며, 만생종은 흑향이 40.6 cm로 가장 짧으며 백진주1호가 56.44 cm로 가장 길었고 평균은 46.98 cm를 나타내었다.
Table 3.
Variation of leaf lengths and widths in three groups of rice varieties in terms of flowering time
Table 3.
Characteristic Min Max Mean SD F p-value
Leaf length ( cm) 53.67*** <0.001
EarlyZ 23.44 47.88 30.96 ±4.55
Medium 21.76 59.10 39.74 ±7.18
Late 40.6 56.44 46.98 ±6.08
Leaf width(cm) 9.12*** <0.001
Early 0.86 1.74 1.27 ±0.17
Medium 1.02 2.54 1.38 ±0.20
Late 1.08 1.8 1.46 ±0.26

ZEarly = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

*P <0.05,

**P < 0.01

***P < 0.001, ns: not significant.

잎몸 너비는 조생종에서 운장벼가 0.86 cm로 가장 좁게 나타 났고 흑가위찰이 1.74 cm로 가장 넓은 잎몸 너비를 가진 것으로 나타났고 평균은 1.27 cm로 관찰되었다. 중생종은 금남벼가 1.02 cm로 최소치를, 성조찰이 2.54 cm로 최대치를 나타냈고 평균은 1.38 cm로 나타났으며, 만생종은 신토흑미가 1.08 cm로 가장 좁게 나타났고 목우가 1.80 cm로 넓게 나타났으며 평균은 1.46 cm로 세 품종군 중에서 가장 넓게 관찰되었다.
출수기에 따른 잎의 길이는 출수기에 따라 유의한 차이가 있는 것으로 나타나서 만생종, 중생종, 조생종 순으로 컸으며, 잎의 너비 또한 출수기에 따라 유의한 차이가 있는 것으로 인정되 어 만생종, 중생종, 조생종 순으로 컸다(Table 3). 만생종에서 잎몸의 길이와 너비의 평균 최고치가 관찰되었으며 이 같은 결과는 엽면적 및 엽신건물중의 증가로 식물체 전체의 광이용율 및 광합성 효율을 극대화시켜 동화물질의 축적에 크게 기여할 것으로 생각된다.
(2)

(2) 잎의 색과 형태

벼 243 품종의 잎의 형태적 특성을 비교 분석하기 위하여 유묘기의 제1엽색, 잎의 녹색 정도 및 안토시아닌 유무, 잎혀의 모양과 색을 조사하였다.
파종 후 3~4엽기에 품종별로 제1엽의 부위별 안토시아닌색소 의 유무를 조사한 결과는 Table 4와 같다. 제1엽색이 녹색을 띄는 품종은 동진벼를 포함하여 235개 품종에서 관찰되었고, 자색을 띄는 품종은 자광찰, 신명흑찰, 흑찰, 풍미1호, 눈보라, 서은, 조생흑찰, 보석측찰, 신토흑미 총 9개 품종으로 조사되었 다. 또한, 자색을 나타내는 부위가 신명흑찰, 신토흑미, 조생흑찰 과 같이 서로 다른 위치에서 부분적으로 나타남을 관찰할 수 있었다(Table 4, Supplementary Table 1).
Table 4.
The investigation of existence of rice varieties showing anthocyanin pigmentation in 1st leaf, leaf blade, and ligule.
Table 4.
Distribution of anthocyanin No. of variety
1st leaf Leaf bade Ligule
Absent 235 239 237
Present 8 4 6

*Variety name were provided in Supplementary Table 1, 2, and 3.

잎과 잎혀에서 안토시아닌색소의 유무를 조사한 결과 주로 흑미로 분류되는 조생흑찰, 보석흑찰, 신토흑미 등에서 나타났고 대부분의 벼 품종에서는 나타나지 않았다(Table 4, Supplementary Table 2, 3).
파종 후 3~4엽기에 품종별로 제1엽색 및 잎의 형태를 조사한 결과는 Table 5와 같다. 벼 품종의 최고분얼기에 잎의 녹색정도 와 잎에 포함된 안토시아닌색소의 유무를 조사하였고, 잎의 녹색 정도는 연녹색, 중간녹색, 진녹색으로 분류하여 관찰하였다. 조 사결과 잎이 연녹색을 나타내는 품종은 고품, 풍미1호로 전체 품종의 0.82%를 차지하였고, 중간녹색은 182개 품종으로 75.31%, 진녹색은 59개 품종으로 23.87%의 비율로 나타났다 (Table 5, Supplementary Table 4).
Table 5.
Number of rice varieties with different green color intensity of leaf blades and with ligule
Table 5.
Green color density No. of variety Shape (Ligule) No. of variety
Pale green 2 Partedness 235
Medium green 182 Sharp 8
Dark green 59

*Variety name were provided in Supplementary Table 4 and 5.

뾰족한 형태의 잎혀를 가진 품종은 만추, 문장, 화랑, 평안, 미품, 흑선찰, 성조찰, 홍진주 총 8개 품종으로 나타났고, 235개 품종은 갈라진 잎혀의 모양으로 나타났다(Table 5, Supplementary Table 4).
(3)

(3) 줄기의 길이 및 분얼수

벼 243개 품종의 줄기의 형태적 특성은 길이 및 분얼수로 나누어 특성 비교를 수행하였다.
이삭의 길이를 제외한 간장과 분얼수를 개화시기별로 분석한 결과는 Table 6과 같다. 조생종은 설백이 62.42 cm로 가장 작았고 자광찰이 105.42 cm로 가장 컸으며 평균 76.86 cm로 나타났고, 중생종은 주안1호가 60.71 cm로 가장 작았고 서농8호 가 84.60 cm로 가장 컸으며 평균 78.20 cm로 3가지 품종군 중에서 가장 크게 나타났다. 만생종은 흑향이 61.16 cm로 가장 작았고 목우가 84.60 cm로 가장 컸으나 평균은 74.32 cm로 가장 작게 나타났다.
Table 6.
Variation of culm lengths and panicles per plants in three rice varieties groups in terms of flowering time.
Table 6.
Characteristic Min Max Mean SD F p-value
Culm length(cm) 1.62ns 0.20
EarlyZ 62.42 105.42 76.86 ±6.62
Medium 60.72 104.60 78.20 ±8.07
Late 61.16 84.60 74.32 ±7.93
Panicles per plant(no.) 3.84* 0.02
Early 6.8 16.0 10.9 ±1.97
Medium 5.6 18.8 11.0 ±2.28
Late 7.4 10.4 8.9 ±1.15

ZEarly = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

*P <0.05

**P < 0.01

***P < 0.001, ns: not significant.

벼의 분얼수는 조생종에서 대찬과 황금보라가 6.8개로 가장 적었고 보석이 16.0개로 가장 많았으며 평균 10.9개로 나타났다. 중생종은 목양이 5.6개로 243개 품종 중 최소치로 나타났고 보석찰이 18.8개로 243개 품종 중 최대치를 나타냈으며 평균 11.0개로 품종군 중 가장 높게 나타났다. 만생종은 목우가 7.4개 로 가장 적었고 서농16이 10.4개로 가장 많았으며 평균은 8.9개 로 나타났다(Table 6).
출수기에 따른 간장의 길이는 출수기 간에 유의성이 나타나지 않았으나, 분얼수의 분석 결과 출수기에 따라 유의한 차이가 있는 것으로 인정되어 중생종이 11개로 가장 많았고 조생종(10.9 개), 만생종(8.9개) 순이었다(Table 6).
(4)

(4) 줄기의 형태와 색

벼 243개 품종의 줄기의 정성적 특성은 출수기 및 황숙기 지엽의 자세, 줄기 굵기 및 자세로 나누어 특성 비교를 수행하였 다.
지엽의 경우 출수기와 황숙기로 나누어 조사를 하였으며, 이삭 을 기준으로 지엽의 각도가 0-20°는 직립형, 35-50°는 반직립형, 50°이상의 각도를 보이는 것은 수평형으로 나누어 구분하였다.
그 결과 출수기에는 222개 품종이 직립형으로 나타났고, 풍미 1호, 두루진미, 운장벼, 오대1호벼 등 36개 품종이 반직립형으로 나타났으며, 수평형을 나타내는 품종은 없었다(Table 7), 황숙기 에는 190개 품종이 직립형, 45개 품종이 반직립형으로 나타났고, 오대1호, 큰눈자미, 청백찰, 홍광, 운두, 대립벼1호, 흑진주, 호반 총 8개 품종이 수평형을 나타내었다(Table 7). 일부 벼 품종을 제외하고는 출수기와 황숙기에 동일한 형태를 띠었으며 대체로 직립형이 많았고, 수평은 극히 일부에 불과하였다. 가장 대표적 인 수평형 품종은 큰눈자미이며, 반직립형 품종은 삼천벼로 나타 났다.
Table 7.
Number of flag leaf angle at heading stage and yellow ripe stage in 243 rice varieties.
Table 7.
Flag leaf attitude No. of variety
Heading stage Yellow ripe stage
Erect 222 190
Semierect 21 45
Medium 0 8

*Variety name were provided in Supplementary Table 6 and 7.

벼의 호숙기 및 성숙기에 줄기 모양을 비교한 결과, 205개 품종이 직립형을 나타냈고, 설백, 남일, 만호, 조아미를 포함한 총 8개 품종이 보통 개형, 29개 품종이 반직립형으로 대부분의 벼 품종이 직립의 형태를 보였다(Table 9). 줄기의 굵기는 24개 품종이 가는 굵기, 164개 품종이 보통 굵기, 55품종이 굵은 줄기로 대부분이 보통의 줄기 굵기를 나타내었다(Table 8).
Table 8.
Number of culm thickness and attitude in 243 rice varieties.
Table 8.
Thickness No. of variety Attitude No. of variety
Thin 24 Erect 205
Medium 164 Semierect 29
Thick 55 Medium 8

*Variety name were provided in Supplementary Table 8 and 9.

Table 9.
Variation of plant height and panicle length in three rice variety groups in terms of flowering time.
Table 9.
Characteristic Min Max Mean SD F p-value
Plant height(cm) 19.55*** <0.001
Early 87.0 135.8 99.8 ±7.95
Medium 91.2 138.1 107.7 ±8.96
Late 85.6 128.1 103.4 ±13.38
Panicle length(cm) 15.18*** <0.001
Early 17.42 25.80 21.51 ±1.68
Medium 15.74 29.64 20.02 ±2.19
Late 16.66 21.70 18.92 ±1.82

ZEarly = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

*P <0.05

**P < 0.01

***P < 0.001, ns: not significant.

(5)

(5) 초장 및 수장

벼 243개 품종을 대상으로 초장 및 수장을 조사하였으며 개화 시기별로 분석한 결과는 Table 9와 같다.
초장의 경우 조생종은 조운이 87.00 cm로 가장 작았고 자광찰 이 135.80 cm로 가장 컸으며 평균 99.80 cm로 나타났고, 중생종 은 호진벼가 91.20 cm로 가장 작았고 서농14호가 138.10 cm로 가장 컸으며 평균 107.70 cm로 3가지 품종군 중에서 가장 크게 나타났다. 만생종은 흑향이 85.60 cm로 가장 작았고 목우가 128.10 cm로 가장 컸으나 평균은 103.40 cm로 나타났다.
벼의 이삭길이는 조생종에서 대진벼가 17.42 cm로 가장 짧았 고 흑진주벼가 25.80 cm로 가장 길었으며 평균 21.57 cm로 품종군 중 가장 높게 나타났다. 중생종은 삼광벼가 15.74 cm로 최소치로 나타났고 목양이 29.64 cm로 243개 품종 중 최대치를 나타냈으며 평균 20.02 cm로 나타났다. 만생종은 슈퍼자미가 16.66 cm로 가장 짧았고 목우가 21.70 cm로 가장 길었으며 평균은 18.92 cm로 나타났다(Table 9).
출수기에 따른 초장의 길이는 출수기에 따라 유의한 차이가 있는 것으로 나타나서 중생종, 만생종, 조생종 순으로 컸으며, 이삭의 길이 또한 출수기에 따라 유의한 차이가 있는 것으로 인정되어 조생종, 중생종, 만생종 순으로 컸다(Table 7).
출수기 및 형태형질 관련 특성
출수기는 조사 개체의 50%가 출수하였을 때를 기준으로 하여 조사하였으며, 이앙 후 일수에 따라 구분하였다. 이앙 후 70일 이전에 출수한 품종은 조생종, 70일에서 90일 사이에 츨수한 품종 은 중생종, 90일 이후에 출수한 품종은 만생종으로 분류하였다(Fig. 1A). 또한, 고품질 벼 품종 재배면적 확대와 품질 고급화 실현을 위해 국내 육성 벼 243 품종을 대상으로 벼 품종별 키, 수형, 이삭 및 종자의 모양과 크기 등 특성을 표시해 농업인들이 올바른 품종 선택으로 고품질 쌀을 안전하게 생산할 수 있도록 정보 제공 및 품종에 대한 구분체계를 확립하여 육종의 기초 자료로 활용하 고자 하였다 (Fig. 1B, Supplementary Figure 1).
Fig. 1.
A, Phenotype of early, medium and late flowering rice varieties at 70 days after transplanting. (A) Boseok, (B) Sinunbong1, (C) Daechan (D) Cheongbaekchal,, (E) Suryeojinmi. B, Morphological traits, plant type, panicle shape and grain size in three representative varieties for early-, medium-, and late-ecotype, respectively, among 243 Korean rice varieties (See Supplementary Fig. 1 for more varieties).
KJBS-48-3-254_F1.gif
다변량 분석에 의한 품종 분류
(1)

(1) 주성분 분석

공시된 243개 벼 품종의 개별적 특성이 객관성이 있게 평가되 어야 하나 단일 변량에 의하여 전체적인 변이의 구조를 파악하기 는 어렵다. 따라서 다변량 분석법을 조사형질보다 적은 수의 주성분으로 요약한 후 여러 변수들의 구조적 관계를 파악하는데 사용하였다(Choi 2011).
벼 243개 품종을 대상으로 출수기, 초장, 간장, 수장, 분얼수, 잎몸너비 및 잎몸길이 총 7가지 형질을 이용하여 통계프로그램 SAS 9.2의 PRINCOMP procedure로 주성분분석을 수행하였 다. 주성분분석으로 고유치와 각 주성분의 기여도를 나타낸 결과 는 Table 10과 같다.
Table 10.
Eigenvalues and cumulative proportion of the first 7 principal components in 243 rice varieties.
Table 10.
Principal component Eigen value Difference Proportion Cumulative
1 2.22194685 0.48697278 0.3174 0.3174
2 1.73497407 0.499865537 0.2479 0.5653
3 1.2351087 0.45826783 0.1764 0.7417
4 0.77684087 0.2008556 0.111 0.8527
5 0.57598527 0.23700687 0.0823 0.935
6 0.3389784 0.22281256 0.0484 0.9834
7 0.11616584 0.0166 1
분석결과 고유치와 기여도에서 각각 제1주성분은 고유치가 2.2로 나타나며 전체 분산 중 31.7%를 설명할 수 있으며, 제2주 성분은 1.73에 24.8%, 제3주성분은 1.24에 17.6%, 제4주성분은 0.78에 11.1%, 제5주성분은 0.58에 8.2%, 제6주성분은 0.34에 4.8%, 제7성분은 0.12에 1.7%로 각각 설명할 수 있었다. 이를 전부 해석하지 않고 이들 중 자료의 변동을 많이 설명하는 몇 개의 주성분만을 선택하는데 일반적으로 고유 값이 1이상, 또는 누적 분산비율이 80% 이상이 되도록 주성분의 수를 결정한다 (RDA 2003b). 본 실험은 고유 값 1이상을 기준으로 3개의 주성분을 사용하여 전체자료 변동의 74%를 설명 할 수 있었는데, 이는 3개의 주성분을 통해 7개 형질에서 측정된 전체 분산의 74%를 설명할 수 있어 벼 243개 품종의 출수기 및 주요 수량구성 요소에 따른 분류가 가능하다는 것을 알 수 있었다. 또한, 이들 주성분이 함축하고 있는 형태적 형질 중 어떤 형질이 이들 3가지 주성분에 요약되어 있는지를 유추하고 시각적으로 분류하여 집단의 경향을 알아보기 위하여 Multibase program을 이용하여 주성분분석을 수행하였다. 농업적으로 중요한 7가지 형태적 형 질 값을 이용하여 품종들의 구조적 관계를 확인한 결과는 Fig. 2와 같다. 제1주성분은 전체변이의 27.1%, 제2주성분은 10%. 제3주성분은 7%를 설명할 수 있었다. 주성분분석 결과 얻어진 243개 품종의 분류적 관계는 품종간의 뚜렷한 집구현상은 관찰 되지 않았지만, 조생종(Red group), 중생종(Yellow group) 및 만생종(Blue group) 크게 3개의 집단으로 구분되었다(Fig. 2B). 이를 바탕으로, 외부형태학적 형질을 조사한 결과 파종 후 개화까 지의 소요일수, 잎의 길이 및 너비에 따라 국내 육성 벼 품종의 분류에 의미가 있는 것으로 확인되었다(Fig. 2A). 또한, 국내 육성 벼 품종을 3개의 집단으로 구분하는 형태 특성 중 출수기가 큰 비중을 차지하는 것을 알 수 있다. 이러한 결과는 타작물의 형태적 특성 즉, 식물크기, 과실의 길이 및 너비 등 차이에 의한 분류 및 유연관계를 분석한 Huh et al.(2008)의 결과처럼 작물의 형태적 형질이 품종간의 분류가 의미가 있음을 보여주었다. 그러 나 정확한 품종군 분류를 위해서는 보다 많은 형태적 형질의 이용과 분자학적 접근이 필요할 것으로 생각된다.
Fig. 2.
Principal component analysis (PCA) on the seven morphological traits of the 243 Korean rice varieties. PC 1 and PC 2 refer to the first and second principal components, respectively.
KJBS-48-3-254_F2.gif
(2)

(2) 군집 분석

군집분석은 주어진 관찰치 중에서 유사한 것들을 몇몇 집단으 로 그룹화하여 각 집단의 성격을 파악함으로써 데이터 전체의 구조에 대한 이해를 돕고자 하는 분석방법으로 주성분분석 자체 로도 정보를 얻을 수 있지만 주성분분석은 차후 분석을 위한 준비단계로 변수의 수를 줄여 그래프를 통한 관찰치의 위치를 시각화하여 집단의 경향을 파악하는데 사용한다(Choi 2011).
벼 243개 품종의 7개 형질을 이용하여 주성분분석을 수행한 후 그래프상의 품종들의 배열로 분류관계를 파악하고자 하였으 며, 전체 변이의 74%를 설명할 수 있는 제3주성분까지의 주성분 점수를 이용하여 군집분석을 실시한 결과는 Fig. 3과 같다.
Fig. 3.
Cluster dendrogram generated by flowering time and phenotypic characters for the 243 rice accessions. *Red: early flowering variety, Yellow: medium flowering variety, Blue: late flowering variety.
KJBS-48-3-254_F3.gif
군집분석을 수행한 결과에 따라서 품종을 분류는 Table 11와 같으며, 0.9에서 4개 군으로 분류되었다. 제І군은 벼 82개 품종이 속하였는데, 그 중 조생종 품종이 60개, 중생종 품종이 22개로 나타났다. 전체적으로 잎몸의 길이 및 너비가 다른 군에 비해 길고 짧지만 많은 분얼수를 가지는 것으로 나타났고, 특이적으로 조생종인 홍광의 출수기가 이앙 후 40일로 월등히 빠른 것으로 나타났다. 제II군은 중생종 품종 114개, 만생종 품종 2개 등 총 116개의 품종이 속하였는데, 다른 군에 비해 잎몸의 길이 및 너비가 긴 특성을 보였다.
Table 11.
Varietal composition of each group classified by the cluster analysis using the first four principal component scores.
Table 11.
Group No. Maturity Variety name
I 82 EarlyZ (60) Geumyeong, Durujinmi, Boseok, Dunnae, Sandeuljinmi, Inwol, Wolbaek, Sambaek, Heukseonchal, Naepungbyeo, Joryeongbyeo, Geurubyeo, Sangsanbyeo, Manna, Sangjuchalbyeo, Sinunbong1, Munjang, Daejin, Seolbaek, Joun, Manho, Josaengheukchal, Saesangju, Jungmo1001, Manan, Jungmo1012, Samcheonbyeo, Handeul, Unjangbyeo, Junghwabyeo, Wonpyeong, Jinbong, Geumseong, Unmi, Joan, Hwadongbyeo, Jopyeong, Jungmo1008, Sangmi, Joami, Jeokjinju, Jungmo1010, Wonpum, Taeseong, Honong, Jungmo1011, Manchu, Goun, Taebong, Ansanbyeo, Wonhae, Jungmo1007, Hyangmibyeo2, Pyeongwon, Cheongbaekchal, Odae1, Undu, Hoban, Heukgawichal, Honggwang
Medium(22) Namcheonbyeo, Hyangmibyeo1, Segyejinmi, Anda, Keunseom, Hanareum2, Geumobyeo1, Dasan2ho, Cheongdam, Gangbaek, Daeanbyeo, Jungsan, Cheonga, Seonong12, Pungmi, Seolhyangchal, Manwol, Dasan, Juanbyeo, Areum, Orae, Dasan1
II 116 Medium(114) Seomyeong, Donghaejinmi, Gopum, Yangjobyeo, Sanggol, Chinnong, Daeyabyeo, Mipum, Geumnambyeo, Deurimi2, Samgwang, Dongjin2, Cheongpungheukchal, Saenuri, Goami2, Goami4, Juan1, Jinbaek, Anmi, Jungmo1006, Jinsumi, Cheongnam, Seojin, Hoan, Daesanbyeo, Dongan, Mihyang, Yeonghaebyeo, Wongwang, Sangok, Heukgwang, Dongwon, Hwamyeongbyeo, Sura, Dongbo, Manpung, Hwabong, Jungan, Wongcheong, Gangchan, Seonong15, Jinsang, Cheongho, Haepyeong, Hojin, Boramchan, Jongnam, Haiami, Suan, Jinbo, Saekyehwa, Jungmo1013, Hanmaeum, Suryeojinmi, Samdeok, Sinbaek, Hwarang, Sinmyeongheukchal, Onnuri, Wonhwang, Deuraechan, Nampyeongbyeo, Junam, Onojjinoijji, Seopyeong, Hwanggeumnodeul, Sampyeong, Jungmo1005, Seolgaeng, Yeongan, Haeoreumi, Sindongjin, Dongjinchal, Baekjinju, Seogan, Nongho, Malgeumi, Cheonghaejinmi, Pyeongan, Dami, Seonong10, Aranghyangchalbyeo, Sujin, Hwasambyeo, Anmi, Hongjinju, Cheongcheongjinmi, Haepyeongchal, Hwaan, Daeripbyeo1, Jinpum, Jungmo1009, Hyangnambyeo, Pungmi1, Hwanambyeo, Daepyeong, Goami3, Manjong, Manmi, Gwangan, Boseokheukchal, Jungmo1004, Migawng, Wonchu, Seoan1, Gancheokbyeo, Heuknam, Seonong9, Seoeun, Seonong6, Boseokchal, Nokwonchal, Seonong8, Saechucheong BIL-3
Late(2) Jungmo1015, Syupeojami
III 31 Medium(24) Namgangbyeo, Keunnunjami, Wonmi, Milkikwin, Hwaseonchalbyeo, Nokyang, Hwasinbyeo, Noreunjachal, Hwasin1, Jungmo1013, Goami, Sobi, Seokjeong, Hwajungbyeo, Geumobyeo2, Hopyeong, Daeripjami, Baekokchal, Dongjin1, Geonganghongmi, Garak, Geuman, Seonong14, Geumtap
Late(7) Seonong16, Sintoheukmi, Heukhyang, Yeonghojinmi, Baekjinju1, Dacheong, Mogwoo
IV 14 Early(6) Hwanggeumbora, Namil, Jeokjinjuchal, Heukjinjubyeo, Hwanggeumchal, Daechan
Medium(8) Hangangchal1ho, Keunnunjami, Nongan, Yunnongchamssal, Mokyang, Nunbora, Jagwangchal, Seongjochal

ZEarly = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties

제І군과 제II군에 속하는 품종이 243개 품종 중 81%의 비율 을 차지하며 제I군에는 조생종 품종이 큰 비율을 차지하고 있으며 제II군에는 중생종 품종이 크게 나타났으며 이것으로 품종을 구분하는 표현형 중 큰 비중을 차지하는 형태적 특성이 출수기임을 알 수 있다. 각 군에 속해 있는 품종을 육종 및 인공교 배에 이용하고자 한다면 출수가 늦은 품종은 파종을 앞당기고 출수가 빠른 품종은 파종을 늦추어 양친의 개화기를 일치시킴으 로써 교잡을 유도할 수 있을 것으로 여겨진다.
제III군은 중생종 24개 품종, 만생종 7개 품종이 속하였으며, III군의 31개 품종은 다른 품종들에 비해 초장과 간장의 길이가 긴 특성을 보이고 있었다. 제IV군은 조생종 품종 6개, 중생종 품종 8개 총 14개 품종이 속하였다. 분얼수는 많지 않으나 초장과 간장의 길이가 월등히 길었으며, 중생종인 성조찰의 잎몸 너비가 2.54cm로 가장 넓게 나타났다. 제III군과 제IV군은 엽면적 및 엽신건물중의 증가로 식물체 전체의 광이용율 및 광합성 효율을 극대화시켜 동화물질의 축적에 크게 기여할 것으로 생각된다.
본 연구에서는 국내 243개 벼 품종에 대하여 형태적 변이의 범위를 이해하고, 이들의 효율적 관리와 유용한 신품종의 발굴을 위한 기초연구로서 형태적 특성조사 결과를 바탕으로 주성분 분석 및 군집 분석을 수행하였다. 출수기를 포함한 초장, 간장, 수장, 분얼수, 잎몸너비, 잎몸길이 총 7가지 특성을 이용하여 군집분석을 실시한 결과, 총 4개의 품종군으로 분류되었다.
제I군과 제II군은 출수기가 중요한 표현형으로 나타났으며 제III군과 제IV군은 특이적으로 우월한 표현형 위주의 분류군으 로 나타났다. 주성분분석과 군집분석 결과 크게 4개 군집, 조생 종, 중생종 및 만생종, 그리고 혼합된 군집 경향으로 분류되어 동일한 결과를 보였다. 품종간의 뚜렷한 군집 현상이 관찰되지 않고, 한 개의 군집내에 숙기에 따른 많은 품종들이 속해 있는 결과는 서로 간에 유사한 표현형을 가지고 있는 유전적 유사도가 높은 교배친이 교배모본으로 이용된 데서 기인한 것이라 생각된 다(Lee et al. 2015).
따라서 본 연구 결과로 고찰해 볼 때 신품종 발굴을 위해서는 육종의 초기세대에서 숙기형태별로 계통을 구분하여 적응성을 평가하는 것이 바람직할 것으로 판단된다.
국내 243개 벼 품종에 대하여 잎과 줄기의 형태적 특성조사 결과를 바탕으로 주성분 분석 및 군집 분석을 수행하였다. 잎몸의 길이 및 너비는 만생종에서 각각 46.98 cm과 1.46 cm로 가장 길고 넓게 나타났으며, 흑미로 분류되는 품종의 제1엽, 본엽, 잎혀에서 안토시아닌 색소를 나타내었다. 잎의 녹색정도는 중간 녹색을 가진 품종의 비율이 높았고 갈라진 잎혀의 모양을 가진 품종이 대부분이었다. 또한, 간장의 길이는 조생종에서 최대치 105.42 cm가 나타났고 중생종에서 최소치 60.72 cm가 나타났으 며, 분얼수는 중생종에서 최소치 5.6개와 최대치 18.8개가 모두 나타났다. 일부 벼 품종을 제외하고는 출수기와 황숙기 지엽 자세가 동일한 형태를 띠었으며 대체로 직립형이 많았고, 수평은 극히 일부에 불과하였다. 출수기를 포함한 정량적인 7가지 특성 을 이용하여 군집분석을 실시한 결과, 총 4개의 품종군으로 분류 되어 주성분분석과 동일한 결과, 조생종, 중생종 및 만생종, 그리 고 혼합된 군집 경향을 보였다. 이와 같은 결과, 신품종 발굴을 위해서는 육종의 초기세대에서 숙기형태별로 계통을 구분하여 적응성을 평가하는 것이 바람직할 것으로 판단된다.
아래와 같은 보충자료는 한국육종학회지 홈페이지 (www.koreabreedjournal.org)에서 확인할 수 있습니다.
Supplementary Table 1. The investigation of existence of rice variety showing anthocyanin pigmentation in 1st leaf.
Supplementary Table 2. The investigation of existence of rice variety showing anthocyanin pigmentation in leaf blade.
Supplementary Table 3. The investigation of existence of rice variety showing anthocyanin pigmentation in ligule.
Supplementary Table 4. Frequency distribution of rice variety showing different green color intensity in leaf blade.
Supplementary Table 5. Frequency distribution of rice variety showing types in ligule.
Supplementary Table 6. Frequency distribution of flag leaf angle in 243 rice varieties at heading stage.
Supplementary Table 7. Frequency distribution of flag leaf angle in 243 rice varieties at yellow ripe stage.
Supplementary Table 8. Frequency distribution of culm thickness in 243 rice varieties.
Supplementary Table 9. Frequency distribution of culm attitude at maturity in 243 rice varieties.
Supplementary Figure 1. Morphological traits, plant type, shape and size of panicle and grain in 243 Korean rice varieties.
본 연구결과는 농림축산식품부의 재원으로 농림수산식품기 술기획평가원의 농생명산업기술개발사업의 지원(313043-03-3 -HD030)을 받아 연구되었으며 이에 대한 감사를 표합니다.
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Classification of Korean Rice Varieties Based on Agro-morphological Traits
Korean. J. Breed. Sci.. 2016;48(3):254-270.   Published online September 30, 2016
DOI: https://doi.org/10.9787/KJBS.2016.48.3.254
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Classification of Korean Rice Varieties Based on Agro-morphological Traits
Korean. J. Breed. Sci.. 2016;48(3):254-270.   Published online September 30, 2016
DOI: https://doi.org/10.9787/KJBS.2016.48.3.254
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Classification of Korean Rice Varieties Based on Agro-morphological Traits
Image Image Image
Fig. 1. A, Phenotype of early, medium and late flowering rice varieties at 70 days after transplanting. (A) Boseok, (B) Sinunbong1, (C) Daechan (D) Cheongbaekchal,, (E) Suryeojinmi. B, Morphological traits, plant type, panicle shape and grain size in three representative varieties for early-, medium-, and late-ecotype, respectively, among 243 Korean rice varieties (See Supplementary Fig. 1 for more varieties).
Fig. 2. Principal component analysis (PCA) on the seven morphological traits of the 243 Korean rice varieties. PC 1 and PC 2 refer to the first and second principal components, respectively.
Fig. 3. Cluster dendrogram generated by flowering time and phenotypic characters for the 243 rice accessions. *Red: early flowering variety, Yellow: medium flowering variety, Blue: late flowering variety.

Fig. 1.

Fig. 2.

Fig. 3.

Classification of Korean Rice Varieties Based on Agro-morphological Traits

List of 243 rice varieties bred in Korea from 1982 to 2013.

Agronomic traits investigated in the morphological analysis of rice varieties.

Characteristics Subsection Stage
Qualitative trait
1st leaf Color Nursery stage
Leaf Intensity of green color Maximum tiller number stage
Distribution of anthocyanin
Ligule Color, Shape Maximum tiller number stage
Flag leaf Attitude Heading stage, Yellow ripe stage
Culm Thickness, Attitude Dough ripe stage, Maturity stage
Quantitative trait
Leaf blade Length, Width (cm) Maximum tiller number stage
Culm Length (cm) Dough ripe stage, Maturity stage
Time of heading Time of heading (Day) 50% heading of rice
Panicles per plant (No.)
Panicle Length (cm) Dough ripe stage, Maturity stage

Variation of leaf lengths and widths in three groups of rice varieties in terms of flowering time

Characteristic Min Max Mean SD F p-value
Leaf length ( cm) 53.67*** <0.001
EarlyZ 23.44 47.88 30.96 ±4.55
Medium 21.76 59.10 39.74 ±7.18
Late 40.6 56.44 46.98 ±6.08
Leaf width(cm) 9.12*** <0.001
Early 0.86 1.74 1.27 ±0.17
Medium 1.02 2.54 1.38 ±0.20
Late 1.08 1.8 1.46 ±0.26

ZEarly = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

*P <0.05,

**P < 0.01

***P < 0.001, ns: not significant.

The investigation of existence of rice varieties showing anthocyanin pigmentation in 1st leaf, leaf blade, and ligule.

Distribution of anthocyanin No. of variety
1st leaf Leaf bade Ligule
Absent 235 239 237
Present 8 4 6

Variety name were provided in Supplementary Table 1, 2, and 3.

Number of rice varieties with different green color intensity of leaf blades and with ligule

Green color density No. of variety Shape (Ligule) No. of variety
Pale green 2 Partedness 235
Medium green 182 Sharp 8
Dark green 59

Variety name were provided in Supplementary Table 4 and 5.

Variation of culm lengths and panicles per plants in three rice varieties groups in terms of flowering time.

Characteristic Min Max Mean SD F p-value
Culm length(cm) 1.62ns 0.20
EarlyZ 62.42 105.42 76.86 ±6.62
Medium 60.72 104.60 78.20 ±8.07
Late 61.16 84.60 74.32 ±7.93
Panicles per plant(no.) 3.84* 0.02
Early 6.8 16.0 10.9 ±1.97
Medium 5.6 18.8 11.0 ±2.28
Late 7.4 10.4 8.9 ±1.15

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

P <0.05

P < 0.01

P < 0.001, ns: not significant.

Number of flag leaf angle at heading stage and yellow ripe stage in 243 rice varieties.

Flag leaf attitude No. of variety
Heading stage Yellow ripe stage
Erect 222 190
Semierect 21 45
Medium 0 8

Variety name were provided in Supplementary Table 6 and 7.

Number of culm thickness and attitude in 243 rice varieties.

Thickness No. of variety Attitude No. of variety
Thin 24 Erect 205
Medium 164 Semierect 29
Thick 55 Medium 8

Variety name were provided in Supplementary Table 8 and 9.

Variation of plant height and panicle length in three rice variety groups in terms of flowering time.

Characteristic Min Max Mean SD F p-value
Plant height(cm) 19.55*** <0.001
Early 87.0 135.8 99.8 ±7.95
Medium 91.2 138.1 107.7 ±8.96
Late 85.6 128.1 103.4 ±13.38
Panicle length(cm) 15.18*** <0.001
Early 17.42 25.80 21.51 ±1.68
Medium 15.74 29.64 20.02 ±2.19
Late 16.66 21.70 18.92 ±1.82

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

P <0.05

P < 0.01

P < 0.001, ns: not significant.

Eigenvalues and cumulative proportion of the first 7 principal components in 243 rice varieties.

Principal component Eigen value Difference Proportion Cumulative
1 2.22194685 0.48697278 0.3174 0.3174
2 1.73497407 0.499865537 0.2479 0.5653
3 1.2351087 0.45826783 0.1764 0.7417
4 0.77684087 0.2008556 0.111 0.8527
5 0.57598527 0.23700687 0.0823 0.935
6 0.3389784 0.22281256 0.0484 0.9834
7 0.11616584 0.0166 1

Varietal composition of each group classified by the cluster analysis using the first four principal component scores.

Group No. Maturity Variety name
I 82 EarlyZ (60) Geumyeong, Durujinmi, Boseok, Dunnae, Sandeuljinmi, Inwol, Wolbaek, Sambaek, Heukseonchal, Naepungbyeo, Joryeongbyeo, Geurubyeo, Sangsanbyeo, Manna, Sangjuchalbyeo, Sinunbong1, Munjang, Daejin, Seolbaek, Joun, Manho, Josaengheukchal, Saesangju, Jungmo1001, Manan, Jungmo1012, Samcheonbyeo, Handeul, Unjangbyeo, Junghwabyeo, Wonpyeong, Jinbong, Geumseong, Unmi, Joan, Hwadongbyeo, Jopyeong, Jungmo1008, Sangmi, Joami, Jeokjinju, Jungmo1010, Wonpum, Taeseong, Honong, Jungmo1011, Manchu, Goun, Taebong, Ansanbyeo, Wonhae, Jungmo1007, Hyangmibyeo2, Pyeongwon, Cheongbaekchal, Odae1, Undu, Hoban, Heukgawichal, Honggwang
Medium(22) Namcheonbyeo, Hyangmibyeo1, Segyejinmi, Anda, Keunseom, Hanareum2, Geumobyeo1, Dasan2ho, Cheongdam, Gangbaek, Daeanbyeo, Jungsan, Cheonga, Seonong12, Pungmi, Seolhyangchal, Manwol, Dasan, Juanbyeo, Areum, Orae, Dasan1
II 116 Medium(114) Seomyeong, Donghaejinmi, Gopum, Yangjobyeo, Sanggol, Chinnong, Daeyabyeo, Mipum, Geumnambyeo, Deurimi2, Samgwang, Dongjin2, Cheongpungheukchal, Saenuri, Goami2, Goami4, Juan1, Jinbaek, Anmi, Jungmo1006, Jinsumi, Cheongnam, Seojin, Hoan, Daesanbyeo, Dongan, Mihyang, Yeonghaebyeo, Wongwang, Sangok, Heukgwang, Dongwon, Hwamyeongbyeo, Sura, Dongbo, Manpung, Hwabong, Jungan, Wongcheong, Gangchan, Seonong15, Jinsang, Cheongho, Haepyeong, Hojin, Boramchan, Jongnam, Haiami, Suan, Jinbo, Saekyehwa, Jungmo1013, Hanmaeum, Suryeojinmi, Samdeok, Sinbaek, Hwarang, Sinmyeongheukchal, Onnuri, Wonhwang, Deuraechan, Nampyeongbyeo, Junam, Onojjinoijji, Seopyeong, Hwanggeumnodeul, Sampyeong, Jungmo1005, Seolgaeng, Yeongan, Haeoreumi, Sindongjin, Dongjinchal, Baekjinju, Seogan, Nongho, Malgeumi, Cheonghaejinmi, Pyeongan, Dami, Seonong10, Aranghyangchalbyeo, Sujin, Hwasambyeo, Anmi, Hongjinju, Cheongcheongjinmi, Haepyeongchal, Hwaan, Daeripbyeo1, Jinpum, Jungmo1009, Hyangnambyeo, Pungmi1, Hwanambyeo, Daepyeong, Goami3, Manjong, Manmi, Gwangan, Boseokheukchal, Jungmo1004, Migawng, Wonchu, Seoan1, Gancheokbyeo, Heuknam, Seonong9, Seoeun, Seonong6, Boseokchal, Nokwonchal, Seonong8, Saechucheong BIL-3
Late(2) Jungmo1015, Syupeojami
III 31 Medium(24) Namgangbyeo, Keunnunjami, Wonmi, Milkikwin, Hwaseonchalbyeo, Nokyang, Hwasinbyeo, Noreunjachal, Hwasin1, Jungmo1013, Goami, Sobi, Seokjeong, Hwajungbyeo, Geumobyeo2, Hopyeong, Daeripjami, Baekokchal, Dongjin1, Geonganghongmi, Garak, Geuman, Seonong14, Geumtap
Late(7) Seonong16, Sintoheukmi, Heukhyang, Yeonghojinmi, Baekjinju1, Dacheong, Mogwoo
IV 14 Early(6) Hwanggeumbora, Namil, Jeokjinjuchal, Heukjinjubyeo, Hwanggeumchal, Daechan
Medium(8) Hangangchal1ho, Keunnunjami, Nongan, Yunnongchamssal, Mokyang, Nunbora, Jagwangchal, Seongjochal

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties

Table 1. List of 243 rice varieties bred in Korea from 1982 to 2013.
Table 2. Agronomic traits investigated in the morphological analysis of rice varieties.
Table 3. Variation of leaf lengths and widths in three groups of rice varieties in terms of flowering time

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

P <0.05,

P < 0.01

P < 0.001, ns: not significant.

Table 4. The investigation of existence of rice varieties showing anthocyanin pigmentation in 1st leaf, leaf blade, and ligule.

Variety name were provided in Supplementary Table 1, 2, and 3.

Table 5. Number of rice varieties with different green color intensity of leaf blades and with ligule

Variety name were provided in Supplementary Table 4 and 5.

Table 6. Variation of culm lengths and panicles per plants in three rice varieties groups in terms of flowering time.

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

P <0.05

P < 0.01

P < 0.001, ns: not significant.

Table 7. Number of flag leaf angle at heading stage and yellow ripe stage in 243 rice varieties.

Variety name were provided in Supplementary Table 6 and 7.

Table 8. Number of culm thickness and attitude in 243 rice varieties.

Variety name were provided in Supplementary Table 8 and 9.

Table 9. Variation of plant height and panicle length in three rice variety groups in terms of flowering time.

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties.

P <0.05

P < 0.01

P < 0.001, ns: not significant.

Table 10. Eigenvalues and cumulative proportion of the first 7 principal components in 243 rice varieties.
Table 11. Varietal composition of each group classified by the cluster analysis using the first four principal component scores.

Early = Early flowering rice varieties; Medium = Medium flowering rice varieties; Late = Late flowering rice varieties

Table 1.

Table 2.

Table 3.

Table 4.

Table 5.

Table 6.

Table 7.

Table 8.

Table 9.

Table 10.

Table 11.

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