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핵심집단 육성을 위한 옥수수 계통의 형태적 특성 연구

Analysis of Morphological Characteristics for a Core Set of 194 Maize Accessions

O Gyeom Kim1, Tak Ki Hong1, Woo Ri Ko1, Su Yeon Woo1, Shi Jun Ma1, Jong Won Park2, Kyu Jin Sa1, Ju Kyong Lee1,*
The Korean Journal of Breeding Science 2016;48(2):133-139.
Published online: May 31, 2016

1 강원대학교 농업생명과학대학 식물자원응용공학과,

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*Corresponding author (jukyonglee@kangwon.ac.kr, +82-33-250-6415, +82-33-244-6410)
• Received: February 17, 2016   • Accepted: April 1, 2016

© The Korean Society of Breeding Science

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  • In order to develop a core set and new corn variety in Korea, we evaluated the morphological characteristics of 194 maize accessions by examining eight quantitative characteristics. On the evaluation of quantitative traits for 194 maize accessions, they showed the morphological variations in tassel length (35.1±5.0 cm), plant height (226.1±33.7 cm), ear height (86.3±22.6 cm), stem diameter (2.3±0.6 cm), leaf width (9.3±1.1 cm), ear length (14.5±2.4 cm), ear row number (14.1±1.9 row), and 100 kernel weight (24.9±4.4 g). The results of principal component analysis (PCA) indicated that the tassel length, plant height, and ear height greatly contributed to positive direction on the first principal component axis. One-hundred kernel weight contributed to negative direction on the second principal component axis. Thus these morphological characteristics, which contributed greatly in the first and second principal components, might be useful for discrimination among 194 maize accessions. In our study, seven accessions, such as IT026357, IT026441, IT027321, IT033271, IT033591, IT033597 and IT124273, particularly were measured high on yield-related traits. Consequently, the 194 maize accessions used in this study could be used as promising materials for maize breeding programs such as development of new hybrid in Korea.
옥수수(Zea mays L.)는 벼, 밀과 함께 세계 3대 식량작물 중 하나로 우리나라를 포함하여 전세계적으로 식용, 간식용, 사료용, 공업용 등 다양한 용도로 이용되고 있다. 우리나라에서 는 주로 식용 옥수수를 재배, 이용하고 있지만, 사료용 옥수수는 대부분 수입에 의존하고 있다. 2014년 국내의 옥수수 수급 동향 을 살펴보면 옥수수 수입량은 약 10,138천 톤으로 많은 양이 수입되고 있으며, 그 중 약 80%(8,104천 톤)가 사료용 옥수수로 이용되고 있다(농림축산식품부 2015. 농림축산식품 주요통계). 더욱이 사료용을 포함한 옥수수의 국내 곡물자급도는 1%에 불과하기 때문에(Son et al. 2015), 국내에서 옥수수의 수입량 감소 및 곡물자급도 향상을 위해서는 국내 환경에 맞는 우수한 옥수수 신품종 개발이 필요하다.
현재 국내 옥수수 품종 육종은 국립식량과학원, 홍천 옥수수연 구소 등의 국가기관의 주도 하에 이루어지고 있으며, 많은 우량 품종들이 육성되었다. 주로 식용 옥수수인 찰옥수수(Park et al. 2007a), 단옥수수(Lee et al. 2014), 초당옥수수(감미옥) 품종 에 관심을 기울이고 있다. 최근 국민의 식생활의 변화로 축산물의 수요가 증가하면서 사료용 옥수수 품종들이 육성되어 이용되고 있지만, 여전히 국내산 옥수수 곡물가격이 수입산과 비교하여 2~3배 이상 높은 가격으로 인하여 경쟁력이 낮은 상황이다(Son et al. 2014, Son et al. 2015). 더욱이 소비자들의 건강에 대한 관심이 급증함에 따라 안토시아닌 색소 등과 같은 기능성 물질이 추가된 옥수수 품종들이 개발되었다(Lee et al. 2011, Park et al. 2007b).
농촌진흥청 국립농업유전자원센터에서는 2008년을 기준으 로 약 7,000 여 개의 옥수수 품종 및 계통들을 수집, 보존하고 있다(RDA 2009). 수집한 작물의 유전자원 보존과 품종 개발을 성공적으로 하기 위해서는 수집된 작물의 유전자원들에 대한 유전적 변이의 다양성을 측정 평가하고 이해하는 것이 매우 중요하다. 최근 수집된 유전자원의 수가 증가하면서 이러한 유전 자원들의 효율적인 관리, 평가, 활용에 많은 제약이 발생하고 있다(Li et al. 2004). 따라서 이러한 문제를 극복하기 위해서 Frankel & Brown (1984)은 전체 유전자원의 유전적 다양성을 포함하는 소수의 핵심집단 육성을 제안하였다. 다양한 수집자원 이나 품종들의 객관적이면서도 종합적인 분류방법인 다변량 분석법(Multivariate analysis)은 형태적 특성에 기초하여 수집 된 집단 및 계통들 내에서 유전변이를 분석하는데 유용한 방법 중 하나이다(Nielsen & Munck 2003). 그 중에서 주성분분석법 (Principal component analysis, PCA)은 형태학적 특성을 이용 하여 수집된 식물 집단의 유전자원 분류에 많이 이용되고 있으며, 효과적이면서도 비교적 용이한 분석법 중 하나이다(Khorasani et al. 2011).
본 연구에서는 옥수수 신품종개발을 위한 자식계통 육성 및 핵심집단 육성을 목적으로 농촌진흥청 국립농업과학원 농업유 전자원센터에서 보존하고 있는 옥수수 유전자원들 중에서 분양 을 받은 194개의 옥수수 유전자원 계통들에 대하여 형태적 특성 조사와 주성분분석을 이용한 형태적 변이를 분석하였다.
공시재료 및 형태적 변이 조사
본 연구에서 분석에 이용한 옥수수 계통들은 2013년도에 국립 농업과학원 농업유전자원센터에서 분양 받은 옥수수 계통들로 서 IT번호는 Table 1에 나타내었다. 이들 분양 받은 옥수수 계통들 중에서 2014년에 총 500계통 옥수수에 대한 형태적 특성을 조사하기 위하여 강원대학교 부속농장에 각 계통 별로 20립씩 종자를 파종하여 20개체를 심었으며, 개체 간 거리는 25 cm, 계통 간 거리는 70 cm로 하여 일반 관행 재배법으로 재배하였다. 형태적 특성 조사는 각 계통별로 10개체씩 생육시기 별로 형태조사를 실시하였으며, 조사한 형질들은 웅수길이 (QN1), 간장(QN2), 착수고(QN3), 간경(QN4), 엽폭(QN5), 이 삭장(QN6), 이삭열수(QN7), 백립중(QN8) 등 총 8개의 양적 형질이다(Table 2). 이들 중에서 계통 고정 여부, 재배 안정성(착 수고) 및 수량성(이삭장, 이삭열수, 백립중) 등을 고려하여, 옥수 수 194계통들을 선발하였다.
Table 1.
List of 194 maize accessions surveyed for morphological character.
Table 1.
Table 2.
Characters used in the morphological analysis of 194 maize accessions.
Table 2.
Abbreviation Character When/how measured Category
Q1 Length of Tassel at flowering stage cm
Q2 Plant Height at maturing stage cm
Q3 Ear Height at maturing stage cm
Q4 Stem Diameter at maturing stage cm
Q5 Leaf Width at maturing stage cm
Q6 Ear Length after harvest cm
Q7 Ear Row Number after harvest row
Q8 100 Kernel Weight after harvest g
Data 통계 분석
옥수수 194계통들에서 측정한 8개의 양적 형질들에 대한 통계 분석은 Microsoft Excel 통계 프로그램 및 NTSYS 2.1 프로그램 을 이용하여 수행하였다. 통계분석은 194개의 옥수수 계통들에 서 조사한 형질들에 대하여 평균 및 편차 등 각 계통들에 대한 형질별 변이 분포를 조사하였다. 194개의 옥수수 계통들에 대한 형태적 변이 수준을 이해하기 8개의 양적 형질을 이용하여 주성 분분석과 상관분석을 실시하였다.
옥수수 계통들의 형태적 변이
본 연구에서 옥수수 194계통들에 대하여 8개의 양적 형질을 조사한 결과는 Table 3에 나타내었다. 그 결과 웅수길이(QN1)는 21.0~49.8 cm의 범위로, 평균 35.1±5.0 cm을 나타내었으며, 간장(QN2)은 150.4~323.4 cm의 범위로, 평균 226.1±33.7 cm 의 값을 나타내었다. 착수고(QN3)의 경우는 평균 86.3±22.6 cm로 최소 40.0 cm에서 최대 171.5 cm의 범위를 나타내었다. 간경(QN4)은 평균 2.3±0.6 cm로 1.3~5.0 cm의 범위를 나타내 었다(Table 3). 일반적으로 내도복성은 옥수수 재배 시 매우 중요한 형질 중의 하나이며, 간경은 도복을 결정하는 중요한 형질들 중 하나인 것으로 확인되었다(Kashiwagi et al. 2008). 이전의 연구 결과, 내도복성을 나타내는 계통들은 착수고율이 50% 내외로 알려져 있으며(Ryu et al. 2001), 본 연구에서 이용 된 옥수수 계통들은 모두 착수고율이 50% 내외였다. 더욱이 간장과 착수고의 길이를 고려했을 때, 간장이 길고 착수고가 높은 품종이 도복이 증가하고 수량감소를 유발한다고 보고되었 다(Aldrich et al. 1986, Kim et al. 1997). 본 연구에 이용된 계통들은 착수고율은 50% 내외였지만, 많은 계통들이 너무 긴 간장과 착수고를 나타내어 도복에 약할 것으로 생각되었다. 엽폭 (QN5)은 평균 9.3±1.1 cm로 최소 5.9 cm에서 최대 12.2 cm의 범위를 나타내었다. 더욱이 엽폭은 엽면적을 구성하는 요인 중에 하나이며(Li et al. 2015), 엽면적은 바이오매스 및 최종 수량에도 간접적으로 관여하는 형질이다(Baute et al. 2015). 따라서 엽폭 역시 옥수수 육종에서 고려할 중요한 형질이다.
Table 3.
Mean, standard deviation and range for 8 quantitative characteristics in 194 maize accessions.
Table 3.
Morphological character Unit Mean ± SD Minimum Maximum
Tassel Length cm 35.1±5.0 21.0 49.8
Plant Height cm 226.1±33.7 150.4 323.4
Ear Height cm 86.3±22.6 40.0 171.5
Stem Diameter cm 2.3±0.6 1.3 5.0
Leaf Width cm 9.3±1.1 5.9 12.2
Ear Length cm 14.5±2.4 8.0 22.0
Ear Row Number row 14.1±1.9 8.0 18.0
100 Kernel Weight g 24.9±4.4 12.0 38.0
수량성과 관련 있는 형질들에서 이삭장(QN6)은 8.0~22.0 cm의 범위로, 14.5±2.4 cm의 평균값을 나타내었다. 이삭열수 (QN7)의 경우는 평균 14.1±1.9 열로, 최소 8.0 열에서 최대 18.0 열의 범위를 나타내었다. 백립중(QN8)은 최소 12.0 g에서 최대 38.0 g의 범위로, 평균 24.9±4.4 g의 값을 나타내었다 (Table 3). 수량성은 이삭과 종실에 관여하는 형질들에 의해서 좌우되며, 본 연구에서는 분석에 이용된 194개의 옥수수 계통들 에 대하여 이삭과 종실에 관련된 3개의 형질들에 대하여 비교적 높은 수치를 나타내는 계통들을 확인하였다. 그 결과, 194개의 옥수수 계통들 중에서 7개의 옥수수 계통(IT026357, IT026441, IT027321, IT033271, IT033591, IT033597, IT124273)들은 조사된 3개의 형질들 중에서 2개 이상의 형질들에서 비교적 높은 특성을 나타내었다. 따라서 이러한 계통들은 앞으로 이삭 및 종실수량이 높은 품종을 육성함에 있어서 유용한 육종소재로 생각되었다.
한편 본 연구에서는 194개의 옥수수 계통들에서 조사된 8개의 양적 형질들에 대하여 5%와 1%의 유의수준에서 상관관계 분석 을 실시한 결과, 5%의 유의수준에서는 총 20개의 형질 조합에서 유의성이 인정되었고, 1%의 유의수준에서는 총 13개의 형질 조합에서 유의한 상관관계를 나타내었다(Table 4). 간장과 착수 고 사이(0.798**)에서 1% 유의수준에서 가장 높은 정의 상관관 계를 나타냈으며, 이삭열수와 백립중 사이(-0.162*)에서만 부의 상관관계를 나타내었다. 더욱이 웅수길이는 간장(0.504**)와 착수고(0.418**) 사이에서 비교적 높은 상관관계를 나타내었다 (Table 4). 더욱이 본 연구에서 가장 높은 상관계수를 나타낸 간장 및 착수고 사이의 매우 밀접한 관계를 확인하였으며, 이들 두 형질은 재식밀도, 도복, 식물체의 건물중 등과 매우 높은 상관관계를 나타내는 것으로 알려져 있으며, 옥수수에서 주요 육종 형질이다(Cai et al. 2012).
Table 4.
Pearson correlation coefficients among 8 quantitative traits in 194 maize accessions.
Table 4.
Q2 Q3 Q4 Q5 Q6 Q7 Q8
Q1 0.504** 0.418** 0.168* 0.234** 0.207** 0.143* 0.027
Q2 0.798** 0.321** 0.227** 0.309** 0.183* 0.183*
Q3 0.291** 0.099 0.280** 0.164* 0.032
Q4 0.314** 0.049 0.142* 0.055
Q5 0.136 0.189** 0.304**
Q6 0.020 0.131
Q7 -0.162*

**Significance at P < 0.01

*Significance at P < 0.05

옥수수 계통들의 주성분분석
본 연구에서는 194개의 옥수수 계통들 사이에서 계통들의 형태적 변이를 식별하기 위하여 8개의 양적 형질을 이용하여 주성분분석을 수행한 결과, 주성분의 기여율을 보면 제 1주성분 에서 제 5주성분까지 전체 변이의 83.683%를 나타내었다(Table 5). 더욱이 고유치(Eigen value)가 1보다 크다는 것은 하나의 주성분이 변수 한 개 이상의 분산을 설명하므로 일반적으로 고유치가 1보다 큰 주성분만을 추출한다(Lee & Park 2003). 본 연구에서는 제 1, 2, 3주성분의 고유치가 1보다 큰 것으로 확인되어 제 1, 2, 3주성분만을 추출하여 이후 과정을 진행하였 다. 더욱이 분석에 이용된 8개의 양적 형질들 중에서 웅수길이 (0.663), 간장(0.880), 착수고(0.805)는 제 1주성분에서 양의 방향에 크게 기여하였다. 반면에 제 2주성분에서는 백립중 (-0.831), 엽폭(-0.516)이 음의 방향으로 비교적 크게 기여하였 으며, 제 3주성분에서는 이삭열수(-0.589)와 엽폭(-0.516)이 음 의 방향으로 비교적 크게 기여하였다(Table 6). 이들 결과를 바탕으로 제 1, 2, 3축의 주성분에서 양과 음의 방향에 크게 기여한 형질들은 분석에 이용한 194개의 옥수수 계통들을 식별 하는데 유용한 형질들인 것으로 생각되었다.
Table 5.
Eigen value estimated from principal component analysis.
Table 5.
Principal component Eigen Value Cumulative Variance
PC1 2.685 33.560
PC2 1.259 49.302
PC3 1.141 63.568
PC4 0.865 74.381
PC5 0.744 83.683
PC6 0.676 92.128
PC7 0.452 97.782
PC8 0.177 100
Table 6.
Eigen vector from the first to third principal components.
Table 6.
Traits PC1 PC2 PC3
Tassel Length 0.663 0.146 0.093
Plant Height 0.880 0.102 0.195
Ear Height 0.805 0.270 0.253
Stem Diameter 0.506 -0.076 -0.467
Leaf Width 0.464 -0.516 -0.516
Ear Length 0.450 -0.144 0.431
Ear Row Number 0.309 0.414 -0.589
100 Kernel Weight 0.227 -0.831 0.117
본 연구에서는 주성분분석 결과를 바탕으로 제 1, 2주성분을 이용하여 194개의 옥수수 계통들에 대하여 scatter diagram을 나타내었다(Fig. 1). 그 결과, 제 1축과 제 2축을 기준으로 4개의 사분면으로 나누어졌으며, 제 1사분면에는 IT027213를 포함하 여 총 42개의 계통들이 위치하였으며, 제 2사분면에는 IT027308 을 포함하여 총 52개의 계통들이 포함되었다. 더욱이 제 3사분면 에는 IT027357를 포함한 총 55개의 계통들이 포함되었다. 마지 막으로 제 4사분면에는 IT028072를 비롯한 총 45개의 계통들이 포함되었다. 이상의 결과에 의하면 형태특성조사에서 엽폭 (QN5), 이삭장(QN6), 이삭열수(QN7) 등의 형질들은 모든 사분 면의 계통들에서 큰 차이를 보이지 않은 반면에, 웅수길이(QN1), 간장(QN2), 착수고(QN3), 간경(QN4)이 비교적 큰 계통들은 제 1, 4사분면에 분포하고 있었으며, 백립중(QN8)이 비교적 높은 계통들은 제 3, 4사분면에 분포하고 있었다(Fig. 1). 이상으 로 본 연구에서 분석에 이용한 194개 옥수수 핵심집단은 앞으로 우량 옥수수 품종 개발에 매우 중요한 육종소재가 될 것으로 생각이 되며, 이들 재료들에 대한 형태적 변이 및 주성분분석의 결과는 이들을 육종소재로 이용함에 있어 계통 육성 및 교배조합 을 예측하는데 유용한 정보를 제공할 것으로 생각된다.
Fig. 1.
Scatter diagram of 194 maize accessions based on principal components I (PC1) and II (PC2). ●: 42 accessions that relatively higher for QN1, QN2, QN3 and QN4 included in first quadrant; ○: 52 accessions included in second quadrant; ■: 52 accessions that relatively higher for QN8 included in third quadrant; □: 45 accessions that relatively higher for QN1, QN2, QN3, QN4 and QN8 included in fourth quadrant.
KJBS-48-2-133_F1.gif
본 연구는 옥수수 핵심집단 및 신품종을 육성하기 위하여 국립농업과학원 농업유전자원센터에서 분양 받은 194개의 옥수 수 계통들에 대하여 총 8개의 양적 형질들을 이용하여 형태적 변이 연구를 수행하였다. 8개의 양적 형질을 조사한 결과, 웅수길 이는 평균 35.1±5.0 cm를, 간장은 평균 226.1±33.7 cm를, 착수 고는 평균 86.3±22.6 cm를 간경은 평균 2.3±0.6 cm를, 엽폭은 평균 9.3±1.1 cm를, 이삭장은 평균 14.5±2.4 cm를, 이삭열수는 평균 14.1±1.9 열을, 백립중은 평균 24.9±4.4 g를 각각 나타내었 다. 주성분분석결과에서는 8개의 양적 형질들 중에서 웅수길이, 간장, 착수고는 제 1주성분에서 양의 방향에 크게 기여하였다. 반면에 제 2주성분에서는 백립중(-0.831)이 음의 방향으로 크게 기여하였다. 따라서 본 연구에서 제 1축 및 제 2축의 주성분에서 양의 방향과 음의 방향에 크게 기여한 형질들은 분석에 이용한 194개의 옥수수 계통들을 식별하는데 유용한 형질들인 것으로 생각되었다. 본 연구에서 이삭과 종실에 관련된 형질들에서 비교적 높은 수치를 나타낸 7개의 계통들(IT026357, IT026441, IT027321, IT033271, IT033591, IT033597, IT124273)은 이삭 및 종실수량이 높은 품종을 육성함에 있어서 유용한 육종소재로 생각된다. 이상으로 본 연구에서 분석에 이용한 194개 옥수수 핵심집단은 앞으로 우량 옥수수 품종 개발에 매우 중요한 육종소재 가 될 것으로 생각이 되며, 이들 계통들에 대한 형태적 변이 및 주성분분석의 결과는 이들을 육종소재로 이용함에 있어 계통 육성 및 교배조합을 예측하는데 유용한 정보를 제공할 것으로 생각된다.
본 논문은 골든씨드프로젝트(식량종자사업단, 213001-04-3- SBA10, PJ009940022015)의 지원에 의하여 연구 되었음.
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Analysis of Morphological Characteristics for a Core Set of 194 Maize Accessions
Korean. J. Breed. Sci.. 2016;48(2):133-139.   Published online June 30, 2016
DOI: https://doi.org/10.9787/KJBS.2016.48.2.133
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Analysis of Morphological Characteristics for a Core Set of 194 Maize Accessions
Korean. J. Breed. Sci.. 2016;48(2):133-139.   Published online June 30, 2016
DOI: https://doi.org/10.9787/KJBS.2016.48.2.133
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Analysis of Morphological Characteristics for a Core Set of 194 Maize Accessions
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Fig. 1. Scatter diagram of 194 maize accessions based on principal components I (PC1) and II (PC2). ●: 42 accessions that relatively higher for QN1, QN2, QN3 and QN4 included in first quadrant; ○: 52 accessions included in second quadrant; ■: 52 accessions that relatively higher for QN8 included in third quadrant; □: 45 accessions that relatively higher for QN1, QN2, QN3, QN4 and QN8 included in fourth quadrant.

Fig. 1.

Analysis of Morphological Characteristics for a Core Set of 194 Maize Accessions

List of 194 maize accessions surveyed for morphological character.

Characters used in the morphological analysis of 194 maize accessions.

Abbreviation Character When/how measured Category
Q1 Length of Tassel at flowering stage cm
Q2 Plant Height at maturing stage cm
Q3 Ear Height at maturing stage cm
Q4 Stem Diameter at maturing stage cm
Q5 Leaf Width at maturing stage cm
Q6 Ear Length after harvest cm
Q7 Ear Row Number after harvest row
Q8 100 Kernel Weight after harvest g

Mean, standard deviation and range for 8 quantitative characteristics in 194 maize accessions.

Morphological character Unit Mean ± SD Minimum Maximum
Tassel Length cm 35.1±5.0 21.0 49.8
Plant Height cm 226.1±33.7 150.4 323.4
Ear Height cm 86.3±22.6 40.0 171.5
Stem Diameter cm 2.3±0.6 1.3 5.0
Leaf Width cm 9.3±1.1 5.9 12.2
Ear Length cm 14.5±2.4 8.0 22.0
Ear Row Number row 14.1±1.9 8.0 18.0
100 Kernel Weight g 24.9±4.4 12.0 38.0

Pearson correlation coefficients among 8 quantitative traits in 194 maize accessions.

Q2 Q3 Q4 Q5 Q6 Q7 Q8
Q1 0.504** 0.418** 0.168* 0.234** 0.207** 0.143* 0.027
Q2 0.798** 0.321** 0.227** 0.309** 0.183* 0.183*
Q3 0.291** 0.099 0.280** 0.164* 0.032
Q4 0.314** 0.049 0.142* 0.055
Q5 0.136 0.189** 0.304**
Q6 0.020 0.131
Q7 -0.162*

Significance at P < 0.01

Significance at P < 0.05

Eigen value estimated from principal component analysis.

Principal component Eigen Value Cumulative Variance
PC1 2.685 33.560
PC2 1.259 49.302
PC3 1.141 63.568
PC4 0.865 74.381
PC5 0.744 83.683
PC6 0.676 92.128
PC7 0.452 97.782
PC8 0.177 100

Eigen vector from the first to third principal components.

Traits PC1 PC2 PC3
Tassel Length 0.663 0.146 0.093
Plant Height 0.880 0.102 0.195
Ear Height 0.805 0.270 0.253
Stem Diameter 0.506 -0.076 -0.467
Leaf Width 0.464 -0.516 -0.516
Ear Length 0.450 -0.144 0.431
Ear Row Number 0.309 0.414 -0.589
100 Kernel Weight 0.227 -0.831 0.117
Table 1. List of 194 maize accessions surveyed for morphological character.
Table 2. Characters used in the morphological analysis of 194 maize accessions.
Table 3. Mean, standard deviation and range for 8 quantitative characteristics in 194 maize accessions.
Table 4. Pearson correlation coefficients among 8 quantitative traits in 194 maize accessions.

Significance at P < 0.01

Significance at P < 0.05

Table 5. Eigen value estimated from principal component analysis.
Table 6. Eigen vector from the first to third principal components.

Table 1.

Table 2.

Table 3.

Table 4.

Table 5.

Table 6.

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