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옥수수 유망 자식계통들에 대한 잡종강세 및 수량관련 형질의 유전분석

Genetic Analyses for Yield Components and Heterosis in Maize Elite Inbred Lines

Jong Yeol Park1,2, Su Yeon Woo1, Kyu Jin Sa1, Ju Kyong Lee1,*
Korean Journal of Breeding Science 2015;47(4):345-356.
Published online: November 30, 2015

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*Corresponding author (jukyonglee@kangwon.ac.kr, Tel: +82-33-250-6415, Fax: +82-33-259-5558) Jong Yeol Park, Su Yeon Woo, and K. J. Sa contributed equally to this work.
• Received: August 13, 2015   • Revised: November 19, 2015   • Accepted: November 19, 2015

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  • Grain yield, agronomic traits, and correlation between genetic distance of 36 F1 hybrids produced by half-diallel crosses using nine maize inbred lines were analyzed. In the analyses of F1 hybrids and their mid-parent heterosis (MPH) for agronomic traits, grain yield showed highest MPH value of 156%. One-hundred kernel weight showed the lowest value of 7%. In addition, when genetic distance based on agronomic traits was estimated, parental inbred lines did not agree with their own pedigree. Therefore it had the limitation to estimate genetic distance using agronomic traits. In this study, 92 SSR markers were used to calculate genetic distance at DNA level. However we did not confirm their own pedigree of nine parental inbred lines. There was no correlation between grain yield and SSR markers. Therefore molecular marker selection was conducted in relation with grain yield by the step-by-step method using 92 SSR markers. The selected nine SSR markers showed a significant positive correlation (r2=0.703**) between grain yield and SSR markers. The correlations between the selected SSR markers and agronomic traits of ear length, ear width, plant height, and ear height were particularly showed positively correlated. The nine SSR markers selected in this study would help predicting heterosis and planning crosses for hybrids in maize breeding programs.
옥수수(Zea mays L.)는 광과 온도의 이용한계와 광합성 능 력이 높은 C4 작물로 고도의 잡종강세를 이용하기 때문에 단 위면적당 수량이 높아 세계 주요 곡물 중 밀(Triticum aesticum L.), 벼(Oryza sativa L.)와 더불어 세계 3대 식량작물로서 오 랜 육종역사를 가지고 있는 대표적인 타가수정 작물의 하나 이다. 옥수수의 품종개발은 미국을 중심으로 이루어져 왔으 며, 1800년대 미국에서 집단선발에 의한 100여 종의 방임수 분품종이 개발되어 재배되었고(Troyer 1999), Shull (1909) 에 의해 단교잡종의 우수성이 밝혀진 이후 옥수수 육종의 획 기적인 전기를 마련하였으나 실용화가 되지 못하였다. 1910 년대 Jones에 의해 복교잡종 이론이 제기되면서 교잡종 옥수 수 연구가 점차 이루어졌지만 1960년대에 비로소 생산력이 우수한 단교잡종이 보급⋅재배되기 시작하였다.
옥수수 육종은 방법에 따라 여러 가지로 발전되어 왔으며 기존의 식물유전자 연관에 대한 연구는 형태학적 형질과 생 리학적 형질을 마커로 이용하는 연구가 주를 이루었으나, 현 재는 복잡한 유전양상 및 유전자 구명을 위한 분자유전학적 기법을 이용한 유전자 지도 작성 등을 직접적으로 육종에 이 용하려는 시도가 이루어지고 있다. 1980년대부터 분자유전학 적 기법의 개발로 다양한 유전자 마커가 개발되었고, 유전자 지도의 작성이 용이하게 되었다. 그리고 육종의 실질적인 적 용 대상인 양적 형질의 구명으로 더욱 효율적인 육종이 기대 된다. 이렇게 작물 육종연구에서 유전자 지도와 QTL 분석을 통하여 질적 및 양적 형질에 관련된 표지마커 개발은 작물육 종에 유용한 정보를 제공할 수 있다(Lombard & Delourme 2001, Tanksley et al. 1996).
옥수수 육종의 가장 큰 목적인 다수성에 초점을 맞춘다면 수량이 높을 가능성이 있는 교배친의 선발은 많은 노력과 시 간, 비용을 절감할 수 있으나, 현재의 우량 교잡종 선발은 포 장시험을 통해 이루어지기 때문에 많은 인적 물적 자원이 소 모된다. 그래서 효과적으로 교배친을 선발하기 위하여 과거에 는 식물 유전자 연관에 대한 연구는 형태학적 형질과 생리학 적 형질들을 마커로 이용하는 기본적 연구가 주를 이루었으 나, 현재는 분자유전학적 기법을 이용한 유전자 지도 작성 등 을 직접적으로 육종에 이용하려는 시도가 이루어지고 있다. 해마다 많은 자식계통이 개발되고 앞에서 많은 연구자들이 형질과 분자유전학적 방법에 의한 잡종강세를 확인하려 하였 으나 효과가 미미하였다. 만약 자식계통만으로 교잡종의 수량 을 예측할 수 있다면 교잡종 선발의 효과를 높일 뿐만 아니라 선발 노력과 시간을 절약할 수 있다. 이렇게 작물 육종연구에 서 유전자지도와 QTL 분석을 통하여 질적 및 양적 형질에 관 련된 표지마커 개발은 작물육종에 유용한 정보를 제공할 수 있다(Ajmone-Marsan et al. 1998, Godstalk et al. 1990, Griffing & Lindstrom 1954, Hallauer 1972, Hallauer et al. 1988, Lamkey et al. 1987, Lee et al. 1989, Lombard & Delourme 2001, Melchinger 1990, Min 2000, Park 1999b, Tanksley et al. 1996).
따라서 본 연구는 미국에서 육성한 9개의 유망 자식계통 (elite inbred lines)을 교배친으로 사용하여 농업형질과 SSRs (Simple Sequence Repeats) 마커를 이용하여 그들의 유전적 거리를 구하고, 자식계통간의 반이면교잡(Half-Diallel Cross) 에 의해 얻어진 36개 F1 교잡종들의 수량과 마커 사이의 관계 를 구명한 후, 옥수수 수량과 관련된 SSR 마커를 선발하여 교 배친 계통들 사이의 잡종강세를 예측하여 교잡종의 평가에 따 른 노력과 시간을 절감하는데 기여하고자 하였다.
본 연구는 옥수수의 종실수량과 분자생물학적 기법의 하나 인 SSR 마커와의 상관을 분석하기 위하여 미국에서 육성한 9개의 유망 자식계통(elite inbred line)을 이용하여 반이면교 잡에 의해 얻어진 36개의 F1 교잡종들에 대한 수량 검정과 분 자유전학적 기법인 SSR 분석을 수행하였다. 9개의 자식계통 및 이들 사이의 F1 교잡종들에 대한 농업형질 관련 특성조사 는 2000년부터 2003년까지 강원도 농업기술원 옥수수연구소 포장에서 수행하였다.
1) 연구재료
본 연구는 미국에서 육성한 A632, ND203, Va85, Mo17, B73, Oh43, C103, B14A, Wf9 등 9개의 유망 자식계통을 사용하여 수행하였다. 연구에 사용된 자식계통들은 Lancaster Sure Crop (LSC) 집단 4개, Iowa Stiff Stalk Synthetic (BSSS) 집단 4개, 기타 1개의 자식계통을 사용하였으며, 사 용된 9 자식계통의 분류 및 근원은 Table 1과 같다.
Table 1.
Maize elite inbred lines used in this study.
Table 1.
Class Line
Lancaster Sure Crop (LSC) Mo17, C103, Oh43, Va85
Iowa Stiff Stalk Synthetic (BSSS) B73, Wf9, B14A, A632
Others ND203
2) 교배조합 구성 및 포장평가
2000년 9개의 자식계통을 이용하여 반이면 교잡에 의해 36개의 F1 교잡종을 생산하여, 2001년부터 2003년까지 3년 간 3지역(강원도 홍천, 양구, 정선)에 파종하여 포장평가를 수 행하였다. 옥수수 파종은 이랑사이 60 cm, 포기사이 30 cm (5,500 주/10a)로 2열씩 난괴법 3반복으로 2립씩 파종하였다. 본엽 3 ~ 4엽기에 1주 1본을 남기고 나머지는 솎아주었으며, 시비수준은 17.4-3-6.9 (N-P2O5-K2O) kg/10a 시비하였고 질 소는 기비 50%, 7 ~ 8엽기에 추비 50%로 분시하였고, 인산 과 가리는 전량 기비로 시용하였다. 그 외의 재배법은 옥수수 표준재배법에 준하여 재배를 하였다. 조사된 농업형질들은 Table 2와 같다. 농업형질 조사요령은 농사시험연구 조사기 준(농촌진흥청)에 준하여 조사하였다.
Table 2.
Morphological traits measured on individual maize plants.
Table 2.
Traits Abbr. Units Remarks
Days to Anthesis DA day Days from sowing to pollen shed
Days to Silking DS day Days from sowing to silk emergence
Plant Height PH cm Height from soil level to top-leaf node at maturity
Ear Height EH cm Height from soil level to top-ear node at maturity
H. turcicum Ht 0~9 0: very strong resistant, 9: very weak susceptible
Corn Borer CB 0~9 0: very strong resistant, 9: very weak susceptible
Stalk Rot SR 0~9 0: very strong resistant, 9: very weak susceptible
Stalk Lodging SL 1~9 1: no lodging, 9: totally lodging
Ear Length EL cm Length of ear
Ear Width EW cm Width of ear
100 kernel Weight HW g Weight of 100 kernels at 15.5% moisture content
Grain Yield GY kg/10a Grain yields per 10a at 15.5% moisture content
3) SSR 분석
(1)

(1) 식물 DNA 추출

식물 DNA 추출은 연구에 사용된 9개 자식계통을 포트 파 종 후 3 ~ 4엽이 출현하였을 때 어린잎을 채취하여 Dellaporta et al. (1983)의 방법에 의해 추출하였다. 본엽 2번째 엽을 소 량(약 0.5 g) 취한 후 1.5 ml tube에 넣은 즉시 액체질소를 넣 어 최대한 곱게 분쇄하여 DNA 추출 buffer (200mM Tris-HCl pH 8.0, 200 mM NaCl, 25 mM EDTA, 0.5% SDS)를 700 ㎕를 첨가하여 잘 혼합한 후 여기에 동량의 chloroform : isoamylalchol (24 : 1) 넣어 약 10분간 흔들어 준 다음 65℃ 항온수조에 20분간 보관한 후 13,000 rpm에서 15분간 원심분리를 한 후 상등액 600 ㎕를 취하여 새로운 tube에 옮긴 후 냉동 보관된 동량의 isopropanol을 넣어 1시 간 동안 -20℃에 보관하였다. 이것을 다시 13,000 rpm에서 15분간 원심분리 후 상등액을 버리고 pellet을 70% 에탄올 (ethyl alchol)로 씻어서 건조 시킨 후 TE buffer (10mM Tris-Cl, 1mM EDTA)을 넣어 충분히 녹인 다음 10 ㎍/㎖ RNase를 첨가 후 37℃에서 약 12시간 처리하여 RNA를 완전 히 제거하였다. 여기에 0.8% agarose gel에서 DNA 유무와 농도를 확인 후 모든 DNA를 동일한 농도(10 ㎍/㎕)로 희석 한 후 4℃에 저장하여 실험을 수행하였다.
(2)

(2) SSR 분석

SSRs 분석에 사용된 primer들은 Research Genetics (Huntsville, AL, USA)에서 구입한 옥수수 92개의 primer들 을 분석하여 본 연구를 수행하였다. PCR 증폭은 Senior et al. (1996)의 방법을 이용하여 수행하였으며, PCR 반응액 20 ㎕에 옥수수 genomic DNA 20 ng, 각 primer 0.2 uM, dNTPs 100 uM, PCR 반응 buffer 1X (10mM Tris-HCl pH 8.8, 1.5 mM MgCl2, 50 mM KCl, 0.1% Triton X 100), 1 unit DNA Taq polymerase를 첨가하여 반응을 시켰다. 반응 시간 및 온도는 94℃에서 5분간 초기 denaturation 시킨 후 94℃에서 1분, 55℃에서 1분간 annealing, 72℃에서 1분 30 초간 extention을 하여 총 35회 반응을 시켰다.
PCR 반응물의 크기와 band 유무를 확인하기 위하여 Ethidium Bromide (EtBr)를 넣은 3% agarose gel에서 6X loading-buffer (40% sucrose, 0.6 mg/ml bromophenol blue)를 혼합하여 5% denaturing polyacrylamide gel에서 전기영동하여 분석하였다. PCR 반응물 6 ㎕를 3X loading-buffer (10mM NaOH, 95% formamide, 0.05% bromophenol blue, 0.05% xylen cyanol FF) 3 ㎕와 혼합한 후 95℃에서 5분간 끓인 다음 7 ㎕를 1시간 동안 예열된 5% denaturing polyacrylamide gel (acrylamide : bisacrylamide; 19 : 1), 7.5 M urea, 1X TBE buffer (0.89M Tris-HCl pH 8.0, 0.89M boric acid, 0.02M EDTA pH 8.0)에서 80 W에서 band 크기에 따라 전 기영동(약 1시간 ~ 1시간 30분)한 후 silver 염색 후 결과를 APC film으로 현상하였다.
4) 통계분석
(1)

(1) 자식계통 간 유전적 거리

교배친 사이의 유전적 거리는 분석결과에 의한 band 유무 (유 1, 무 0)를 표시한 후 NTSYS-pc.V.2.1 프로그램(Rohlf 1998)을 이용하여 총 표본의 크기에 대한 동형의 비율인 단 순동형계수(simple matching coefficient)를 구하여 유전적 거리(1 - 단순동형계수)를 구한 후 이를 clustering 하였고, dendrogram 결과에서 계통간 유사도를 조사한 후 이미 알려 진 계보도와 비교하였다.
(2)

(2) 조합능력 및 잡종강세

연구에 사용된 9개 자식계통 간에 반이면 교잡을 실시하여 자식계통과 36개의 F1 교잡종에 나타난 수량을 분산분석을 실시한 후 조합능력을 평가하였다. 그리고 조사된 농업형질에 대하여 자식계통과 교잡종을 이용하여 양친의 평균에 대한 잡종강세(mid-parents heterosis, MPH)를 구하였다. 잡종강 세(%)는 (F1-MP)/MP × 100의 식으로 구하였다(Hayes et al. 1955).
(3)

(3) 상관분석

옥수수 F1의 교배조합별 농업형질들에 대한 상관을 분석하 였고, 또 옥수수 종실수량과 SSR 분석결과를 이용한 유전적 거리와의 상관도 분석하였다. 상관분석을 위하여 SAS 및 EXCEL 통계프로그램을 이용하였다. 그리고 분석에 이용된 92개의 SSR primer들 중에서 종실수량과 정의 상관만을 나 타내는 마커를 선발하기 위하여 step-wise regression을 이용 한 step-by-step 방식으로 92개의 마커들에 대하여 각각 한 개씩 종실수량과의 상관분석을 실시하였다.
1) 교배친의 농업적 특성
본 연구에 사용된 교배친은 Lancaster Sure Crop (LSC) 집단 4개, Iowa Stiff Stalk Synthetic (BSSS) 집단 4개, 기타 1개 총 9개 자식계통으로 구성되어 있으며, 각 자식계통들의 생육 및 수량특성은 Table 3과 같다. 화분비산기는 Oh43과 C103이 가장 빨랐고, 대부분 계통이 82일에서 88일이었으며, 출사일수는 85일에서 91일로 화분비산기와 같은 경향을 보였 고 뚜렷한 숙기 차이는 없었다. B14A는 출사기와 화분비산 기 차이(Anthesis-Silking Interval, ASI)가 5일로 다른 계통 들보다 큰 경향을 보였다. 간장은 자식계통들 간의 뚜렷한 차 이를 보였으며, Va85가 209 cm로 가장 컸고, C103이 141 cm로 가장 작았으며, 착수고도 간장과 비슷한 경향을 보였다. 수량성은 Mo17이 457 kg/10a로 가장 많았으며, B14A가 246 kg/10a로 가장 낮았고, Mo17과 C103이 400 kg/10a 이 상의 높은 수량성을 보였으며, B14A와 ND203이 300 kg/10a 이하의 낮은 수량성을 보였다. 이삭길이는 Va85가 16.2 cm로 가장 길었고, A632가 9.8 cm로 가장 짧았다. 그 리고 100립중은 Oh43이 24.4 g으로 가장 낮았고, Va85가 39.7 g으로 가장 높았으며 계통간의 차이가 매우 컸다.
Table 3.
Means of 8 agronomic traits in 9 maize inbred lines.
Table 3.
Inbred line DA (day) DS (day) PH (cm) EH (cm) EL (cm) EW (cm) HW (g) GY (kg/10a)
A632 88 90 148 67 9.8 3.1 27.1 395
ND203 88 90 199 101 10.8 2.8 28.9 281
Mo17 86 90 153 66 13.8 3.4 32.2 457
Oh43 82 85 166 70 16.9 3.4 24.4 385
C103 82 85 141 56 13.2 3.1 29.1 423
B14A 86 91 159 69 12.2 3.4 27.1 246
Va85 86 89 209 105 16.2 3.8 39.7 389
Wf9 86 89 203 109 11.8 3.3 30.3 347
B73 88 90 159 65 11.2 3.6 33.4 334
2) 교잡종의 주요 농업형질과 상관
9개의 자식계통을 교배친으로 반이면 교잡을 통해 얻어진 36개의 F1 교배조합들에 대한 생육 및 수량특성을 조사하였 다(Table 4). 화분비산기는 72일에서 77일이었고, 출사일수 는 75일에서 79일로 자식계통과 마찬가지로 뚜렷한 숙기 차 이는 없었다. 도복은 ND203/Wf9, Va85/Wf9, Va85/B73 등 3교잡계가 가장 약하였고, 나머지 33개 교잡계는 전반적으로 도복에 강하였다. 내병성 정도는 저항성과 이병성을 모두 나 타내어 교잡계간의 차이가 있었고, 내충성도 또한 비슷한 경 향을 보였다. 간장은 219 cm에서 272 cm로 교잡계간 큰 차 이를 보였고, Wf9/B73이 가장 컸고, Oh43/B14A가 가장 작 았다. 착수고도 간장과 비슷한 경향을 보였다. 수량성은 Mo17/B14A가 1,097 kg/10a로 가장 높았고, A632/B14A가 721 kg/10a로 가장 낮았다.
Table 4.
Means of 12 agronomic traits in 36 cross combinations.
Table 4.
Cross Combination DA (day) DS (day) SL (1-9) Ht (0-9) SR (0-9) CB (0- PH (cm) EH (cm) EL (cm) EW (cm) HW (g) GY (kg/10a)
A632/ND203 73 76 2 4 3 3 226 112 16.3 4.1 32.4 887
A632/Mo17 73 76 1 0 0 0 247 130 17.7 4.2 31.5 955
A632/Oh43 72 76 1 4 4 4 229 106 15.8 3.9 31.5 845
A632/C103 73 76 1 0 0 0 226 105 16.2 4.3 26.2 808
A632/B14A 73 76 1 6 4 4 233 111 14.8 4.1 32.6 721
A632/Va85 74 76 3 5 4 4 242 124 18.3 4.5 32.1 1,015
A632/Wf9 73 76 1 6 4 4 247 125 16.9 4.5 31.9 977
A632/B73 75 78 2 7 4 4 228 107 15.1 4.4 30.9 844
ND203/Mo17 73 76 1 0 0 0 239 132 17.6 4.2 31.1 849
ND203/Oh43 73 76 1 0 0 0 246 127 18.9 4.3 33.4 824
ND203/C103 73 76 1 0 0 0 233 117 18.6 4.2 32.3 859
ND203/B14A 73 77 4 5 4 5 251 134 18.1 4.4 34.2 944
ND203/Va85 77 79 1 0 0 0 241 122 17.5 4.4 31.1 968
ND203/Wf9 73 76 6 6 4 4 255 136 16.4 4.4 33.9 862
ND203/B73 72 76 1 2 2 1 243 128 17.0 4.3 33.0 888
Mo17/Oh43 72 75 1 4 4 4 232 109 18.1 4.3 34.3 889
Mo17/C103 73 76 1 0 0 0 229 106 17.5 4.1 27.8 802
Mo17/B14A 73 76 1 5 4 4 248 122 18.1 4.6 36.4 1,097
Mo17/Va85 73 76 3 4 4 5 247 119 19.7 4.5 34.3 1,080
Mo17/Wf9 72 75 2 6 4 4 263 133 17.6 4.5 35.0 1,068
Mo17/B73 74 77 2 6 4 5 225 104 18.2 4.3 33.4 805
Oh43/C103 74 77 1 2 1 2 224 108 17.1 4.1 29.1 816
Oh43/B14A 74 77 1 5 4 4 219 103 16.6 4.0 31.3 757
Oh43/Va85 72 75 1 4 4 4 244 115 20.0 4.4 36.1 995
Oh43/Wf9 75 76 1 0 0 0 259 134 16.5 4.2 28.7 864
Oh43/B73 73 76 1 5 4 4 237 112 18.7 4.3 34.4 876
C103/B14A 74 76 1 3 3 3 238 110 17.5 4.4 29.3 896
C103/Va85 73 76 3 5 4 4 245 118 18.2 4.4 29.4 889
C103/Wf9 73 76 2 7 4 4 266 134 16.8 4.6 31.1 975
C103/B73 74 77 1 5 4 5 236 108 17.5 4.4 29.4 759
B14A/Va85 74 77 3 4 3 4 255 120 18.7 4.6 34.4 1,045
B14A/Wf9 73 77 2 3 3 2 258 139 17.8 4.5 35.5 978
B14A/B73 75 77 1 6 4 5 228 103 17.1 4.3 31.2 938
Va85/Wf9 73 75 5 4 3 3 269 143 18.4 4.5 31.0 1,017
Va85/B73 74 76 5 4 4 4 240 112 19.1 4.4 32.8 1,064
Wf9/B73 73 76 1 8 4 3 272 142 16.3 4.6 32.5 1,012
전반적으로 Mo17, B73, Va85와 교잡한 교잡계가 수량이 높아 일반조합능력이 우수한 계통으로 추정할 수 있었고, A632, C103, Oh43과 교잡된 교잡계가 일반조합능력이 낮은 것으로 추정된다. 이것은 Min (2000)이 Mo17이 포함된 교잡 종은 수량성이 높았고, Oh43으로 구성된 교배조합은 수량이 낮았다는 보고와 비슷한 경향을 보였다. Lancaster Sure Crop (LSC) 집단에서 유래된 계통이 Iowa Stiff Stalk Synthetic (BSSS) 집단에서 유래된 계통보다 조합능력이 우수한 결과를 보인 것도 Min (2000)의 결과와 비슷하였다. 이삭길이는 Oh43/Va85가 20.0cm로 가장 길었으며, A632/B14A가 14.8 cm로 가장 짧았다. 그리고 100립중은 A632/C103이 26.2 g 으로 가장 낮았고, Mo17/B14A가 36.4 g으로 가장 높았으나, 자식계통보다 변이가 적었다.
8개의 형질에 대하여 36개 교배조합의 F1과 교배친의 잡종 강세 평균 및 범위를 Table 5에 나타내었다. F1에 대한 잡종 강세 정도는 종실수량에서 156%로 가장 컸고, 다음으로 착수 고가 54%, 그리고 간장이 43%의 순으로 나타났고, 100립중 이 가장 작은 7%로 나타났다. 이것은 이삭장에서 잡종강세 정도가 가장 높았다는 Park (1999a)과는 조금 다른 결과를 보였으나, 나머지 형질은 본 연구의 결과와 비슷하였다. Park (1999b)의 잡종강세 정도가 종실수량에서 3배로 가장 높았 고, 착수고와 간장이 다음으로 높았다는 것과 일치하는 결과 를 보였다. 화분비산기와 출사일수의 부의 방향으로 잡종강세 정도를 보였고 이것은 잡종강세가 클수록 숙기가 빨라지는 것을 의미하는 것으로 Park (1999b)과 같은 결과를 보였다.
Table 5.
Means and ranges of F1 hybrids and their mid-parent heterosis (MPH) in 36 cross combinations.
Table 5.
Trait F1 MPH (%)
Means Ranges Means Ranges
DA 73 72~77 -14 -17~-10
DS 76 75~79 -14 -17~-9
PH 242 219~272 43 18~64
EH 120 103~143 54 18~95
EL 17.5 14.8~20.0 26 7~44
EW 4.4 3.9~5.5 21 10~52
HW 32.1 26.2~36.4 7 -15~25
GY 913 721~1,097 156 82~258

※MPH (Mid-Parent Heterosis) = (F1-((P1+P2)/2))/(P1+P2)/2)×100

더욱이 36개의 교배조합의 F1과 그들의 교배친 평균에 대 한 잡종강세(MPH)에서 종실수량과 다른 7개의 형질간의 상 관분석 결과, 교잡종 자체에서는 화분비산기와 출사기를 제외 한 모든 형질이 종실수량과 유의성이 있었다(PH: 0.63**; EH: 0.48**; EL: 0.49**; EW: 0.43**; HW: 0.51**). 교배 친 평균에 대한 잡종강세(MPH)에서 종실수량과 형질간의 상 관은 화분비산기와 출사기에서 부의 방향으로 높았는데(DA: -0.48**; DS: -0.44**), 이것은 숙기가 옥수수 수량에 미치는 영향이 부의 방향으로 기여도가 있어 만생종일수록 수량이 높다는 것을 보여주고 있다. Park (1999b)의 100립중이 수량 의 잡종강세 기여도가 가장 높을 것이라는 결과와 달리 본 연 구에서는 수량구성요소 중 이삭길이가 수량에 가장 큰 영향 을 미치는 것으로 나타났고, 다음으로 100립중으로 나타났다.
Fig. 1은 종실중에서 교배친 평균수량, 교잡종 수량 및 교 잡종 평균에 의한 잡종강세 간의 상관을 나타내었다. 그 결과 교배친의 평균수량과 교잡종 종실수량과는 상관이 없었고, 교 잡종 수량과 교잡종 평균에 의한 잡종강세는 정의 상관을, 교 배친 평균수량과 교잡종 평균에 의한 잡종강세는 부의 상관 을 보였다. 이것은 교배친의 평균 종실수량이 높을수록 두 교 배친에 의해 구성된 교잡종의 잡종강세 정도는 낮아지는 것 을 의미하였다.
Fig. 1.
Correlations between mid-parent heterosis (MPH), average yields of parents and grain yields (GY) in F1 hybrids of 36 cross combinations.
KJBS-47-345_F1.gif
3) 수량구성요소에 대한 조합능력 검정
수량구성요소인 종실수량, 100립중, 이삭길이, 이삭폭에 대 한 일반조합능력(GCA: general combining ability) 및 특정 조합능력(SCA: specific combining ability)을 분석하였다. 그 결과 종실수량에서 일반조합능력이 가장 높은 계통은 Va85로 정의 방향으로 109.9를 보였고, 가장 낮은 계통은 C103으로 부의 방향으로 71.4를 보였다. 수량적인 측면에서 이상적인 조합은 Va85를 포함하는 조합이라 추정할 수 있었 다. 종실수량에 대한 특정조합능력이 가장 높은 조합은 BSSS 그룹과 LSC 그룹의 교잡이면서 과거에 수량이 높다고 알려 진 Mo17/B14A가 139.3으로 가장 높았고, 가장 낮은 조합은 BSSS 그룹간의 교잡인 A632/B14A 교잡종이 -166.3으로 낮 았다. 이 결과로 일반조합능력이 가장 높은 Va85의 다른 계 통들과의 특정조합능력이 높지 않은 결과를 보여 일반조합능 력이 높은 계통이 반드시 최고의 수량을 나타내지는 않는다 는 것을 보여주고 있다(Table 6). 그러나 종실수량과는 달리 나머지 3개의 형질(100립중, 이삭길이, 이삭폭)은 품종 간 조 합능력의 효과가 미미하였다. 결론적으로 100립중과 이삭특 성은 계통간 교잡에서 큰 조합능력을 기대하기 어려울 것으 로 생각되어 품종 육성 과정에서 교배조합 구성에 있어 종실 수량만이 계통 간의 조합능력을 기대할 수 있었다.
Table 6.
GCA and SCA effects of grain yields calculated in half-diallel cross of 9 maize inbred lines.
Table 6.
SCA effects GCA effects
ND203 Mo17 Oh43 C103 B14A Va85 Wf9 B73
A632 41.9 43.6 30.6 2.4 -166.3 28.1 35.9 -16.1 -36
ND203 -66.6 5.4 49.3 52.6 -23 -83.3 23.7 -31.9
Mo17 4.1 -74 139.3 22.7 56.4 -125.6 34.4
Oh43 37 -103.7 34.7 -50.6 42.4 -62.6
C103 44.1 -62.4 69.3 -65.7 -71.4
B14A 11.9 -9.4 31.6 10.3
Va85 -70 58 109.9
Wf9 51.7 64.1
B73 -16.9
4) 양친의 유전적 거리와 농업형질과의 상관
(1)

(1) 농업형질에 의한 표현형적 거리와 농업형질과의 상관

시험포장에서 조사한 농업형질로부터 얻어진 교배친들의 표현형적 거리를 계산하여 표현형적 차이로부터 연관 분석하 여 diagram을 나타내었다(Fig. 3a). 그 결과 BSSS 그룹과 LSC 그룹 중 Wf9와 Mo17만이 그룹에서 이탈하였다. Fig. 3a에서 크게 3개의 그룹으로 구분이 되었다. 첫 번째는 LSC 그룹인 Va85 계통이 포함되었고, 두 번째는 LSC 그룹인 Oh43과 C103 계통이 포함되었고, 그리고 세 번째에는 나머 지 6계통들이 포함되어 있었다. 본 연구결과에서 농업형질을 기초로 한 표현형적 거리와 실제 육성된 계보와는 차이가 있 었으며, 농업형질로만 유전적 거리를 측정하는 것은 한계가 있는 것으로 보였다.
농업형질을 기초로 한 표현형적 거리와 주요 농업형질의 잡종강세 정도와의 상관을 분석하였으며, 대부분의 형질에서 상관이 없었으나 교잡종 자체의 이삭길이는 정의 상관을 보 였고(r2=0.43**), 평균에 대한 잡종강세와는 이삭길이에서 부 의 상관을 보였고(r2=-0.37*) 나머지 형질들은 유의성이 없었 다. 이것은 Park (1999b)의 종실중과 상관이 있다는 보고와 차이가 있었으나 나머지 형질과는 비슷한 경향을 보였다. 농 업형질의 표현형적 거리에 의한 옥수수 수량의 잡종강세 예 측은 어려울 것으로 생각되어 다른 방법에 의한 유전적 거리 측정이 필요한 것으로 생각되었다.
(2)

(2) SSR 분석에 의한 유전적 거리와 농업형질과의 상관

9개의 자식계통에 대한 SSR 분석에서 92개의 SSR primer 들을 이용하여 총 349개의 다형화 밴드를 얻었으며, primer 조합당 2 ~ 7개의 다형화 밴드를 나타내어 평균 다형화 밴드 는 3.71개였다(Table 7, Fig. 2). SSR 분석을 통한 얻어진 349개의 다양화 밴드를 이용한 9개 자식계통들 간의 유전적 거리를 계산하여, 이를 근거로 계통유연관계 분석을 실시하여 diagram을 나타내었다(Fig. 3b). 그 결과 BSSS그룹과 LSC 그룹간의 육성 계보에 의한 유연관계를 명확하게 나타내지 못하였다. 더욱이 표현형적 거리와 SSR 마커에 의한 유전적 거리에서도 유의성을 찾을 수 없었다(r2=0.0514ns).
Fig. 2.
An example of an SSR profile of 9 maize inbred lines.
KJBS-47-345_F2.gif
Fig. 3.
Cluster diagram among the 9 maize inbred lines based on average phenetic distance (a), genetic distance of 92 SSR primers (b), genetic distance of 9 SSR primers (c).
KJBS-47-345_F3.gif
Table 7.
Number of allelic bands and PIC values of the SSR loci among 9 maize inbred lines.
Table 7.
Locus No. of allele PICz Locus No. of allele PIC Locus No. of allele PIC Locus No. of allele PIC
phi014 2 0.2 bnlg589 4 0.67 phi053 2 0.8 phi075 2 0.21
phi016 2 0.49 bnlg439 6 0.75 phi057 3 0.37 phi081 4 0.2
phi027 2 0.49 phi011 2 0.64 phi059 2 0.44 phi091 5 0.33
phi031 3 0.53 phi019 6 0.75 phi076 6 0.27 phi092 4 0.38
phi037 3 0.73 phi006 7 0.79 phi077 7 0.59 phi102 4 0.51
phi047 3 0.68 phi037 3 0.73 phi078 4 0.2 phi040 2 0.51
phi073 2 0.35 phi086 4 0.22 phi079 3 0.77 phi058 3 0.16
phi083 2 0.49 phi021 5 0.8 phi080 4 0.56 phi061 4 0.36
phi087 2 0.49 phi026 5 0.77 phi082 3 0.53 phi062 4 0.25
phi090 2 0.51 phi032 2 0.49 phi084 2 0.69 phi085 6 0.85
phi119 4 0.72 phi098 4 0.49 phi096 3 0.09 phi093 3 0.09
phi127 3 0.57 phi113 6 0.7 phi097 2 0.44 phi117 3 0.38
bnlg278 4 0.62 phi125 3 0.64 phi028 5 0.81 phi118 6 0.35
bnlg386 2 0.78 phi033 3 0.57 phi039 4 0.81 phi049 2 0.1
bnlg161 6 0.8 phi034 3 0.57 phi112 4 0.52 phi116 3 0.44
bnlg108 4 0.73 phi035 2 0.44 phi114 2 0.21 phi121 4 0.49
bnlg125 6 0.64 phi038 2 0.49 phi054 5 0.79 phi123 6 0.56
bnlg166 5 0.69 phi041 3 0.57 phi056 7 0.69 phi068 5 0.8
bnlg420 6 0.51 phi044 2 0.35 phi063 3 0.57 phi126 5 0.8
bnlg210 5 0.77 phi045 2 0.78 phi064 3 0.68 phi129 4 0.6
bnlg594 4 0.84 phi089 6 0.7 phi065 7 0.51 2-7 0.09-0.85
bnlg640 4 0.81 phi048 3 0.53 phi066 2 0.44 Means 3.71 0.55
bnlg155 5 0.8 phi050 3 0.73 phi067 4 0.36
bnlg398 2 0.54 phi051 4 0.64 phi072 2 0.69

zPIC : Polymorphic Information Content.

5) 교잡종 종실중과 유전적 거리와의 상관
Hallauer et al. (1988)은 옥수수 육종에서 조합능력이 우 수한 자식계통을 선발하고자 많은 시간과 노력을 소비하고 있으며, 그 이유는 유전자원에서 유래된 자식계통의 포장 수 량 검정 과정에서 수많은 조합으로 인해 광대한 포장시험을 거쳐야 한다고 하였다. 그래서 많은 연구자들이 간편하게 선 발할 수 있는 방법을 찾으려고 노력하였고, 초기에는 지리적 거리(Griffing & Lindstrom 1954, Hallauer 1972, Moll et al. 1965), 형태적 거리(Smith & Smith 1989), 염색체 크기 나 위치(Moll et al. 1972), 동위효소(Lamkey et al. 1987, Price et al. 1986) 등과 수량의 연관을 찾으려 하였으나 큰 효과를 보지 못하였다. 그 이후 새로운 유전적 측정방법의 하 나인 분자마커에 의한 유전적 거리와 수량과의 상관을 구명 을 위한 연구가 진행되고 있다. Melchinger et al. (1990)과 Godstalk et al. (1990)은 RFLPs의 유전적 거리 측정의 유용 성을 분석하였으나 옥수수 자식계통의 계보를 밝힐 수는 있 으나, 수량과의 상관은 예측하기에 무리가 있다고 하였다. Lee et al. (1989)은 RFLPs의 유용성 평가에서 수량 특수조 합능력과 분자유전학적 거리와 유의성이 있다고 하였다. 더욱 이 Song et al. (1998)은 콩에서 SSR 기법에 의한 분자마커 가 질적 형질에는 관련이 있으나 양적 형질과의 관계를 밝히 지 못하였다. RFLPs와 잡종강세 발현에 따른 상관분석을 위 해 Godstalk et al. (1990)의 연구에서 이형집단의 분류에는 가능하나 유전적 거리와 교잡종 특성발현과는 확실한 관계를 밝히지 못하였다. Park (1999b)은 RAPDs를 이용하여 옥수 수 종실수량성과 정의 상관이 있는 primer 4개, 부의 상관이 있는 primer 3개를 선발하였다. Ajmone-Marsan et al. (1998)은 옥수수에서 유전적 마커에 의한 거리와 수량과는 유의성이 있었으나, 너무나 낮아 수량성 예측에 문제가 있다 고 하였고, Reif et al. (2003)은 열대옥수수를 가지고 SSRs 마커와 옥수수 수량과의 연관을 분석한 결과 수량과 유의성 이 매우 높다고 하였다.
본 연구에서는 이러한 연구 결과를 토대로 SSR 분석에 의 한 유전적 거리와 옥수수 수량과의 상관을 찾기 위해 92개의 SSR primer들을 이용한 옥수수 수량과 유전적 거리를 분석 하였으며(Fig. 3b), 앞선 대부분의 연구자들이 언급한 것처럼 옥수수 종실수량과 SSR 분석을 통한 유전적 거리와는 유의 성이 없었다(r2=0.0497ns). 더욱이 SSR 분석을 기초로 한 유 전적 거리와 주요 농업형질의 잡종강세 정도의 상관은 표현 형적 거리와 마찬가지로 대부분의 형질에서 상관이 없었으나 교잡종 자체의 이삭길이만이 정의 상관을 보였고(r2=0.46**), 평균에 대한 잡종강세와는 모든 형질에서 유의성이 없었다. 이것은 Park (1999b)의 RAPD에서 수량과 정의 상관이 있다 는 보고와 다른 결과를 나타내었다.
따라서 본 연구는 모든 마커들이 종실수량에 정의 방향의 상관과 부의 방향 상관이 공존하기 때문에 모든 마커를 이용 한 수량과 유전적 거리에 의한 상관분석은 의미가 없다는 가 정하에 이미 분석이 완료된 92개의 SSR primer들을 가지고 step-by-step 방식으로 옥수수 종실수량에 영향을 미치는 SSR primer를 선발하고자 하였다. 그 결과 옥수수 종실수량 과 정의 상관을 보이는 9개의 primer 조합(phi051, phi057, phi80, phi112, phi113, phi125, phi127, bnlg210, bnlg398) 을 선발하였다.
선발된 9개의 SSR primer들을 이용한 유전적 거리를 근거 로 diagram을 나타내었다(Fig. 3c). 그 결과 옥수수 계통들의 계보와 선발된 SSR을 이용한 유전적 거리는 앞에서 언급한 것처럼 어떤 연관성을 찾을 수 없었다. 이것은 그룹 내의 옥 수수 계보는 유전적으로 가까워야 한다는 가정에서 출발한 오류인 것으로 추정된다. 이번 연구의 궁극적 목적은 옥수수 종실수량과 유전적 거리와 상관이 있는 분자마커 선발이기 때문에 종실수량과 선발된 9개의 SSR primer를 이용한 유전 적 거리와의 상관은 고도의 정의 상관(r2=0.703**)을 보였다. 이 같은 결과는 기존의 결과와 다른 결과이며, Park (1999b) 가 수량과 정의상관을 보이는 primer 조합을 선발하면 잡종 강세 예측을 더욱 높일 수 있다고 한 결과와 일치하였다.
선발된 SSR primer를 이용한 유전적 거리와 교잡종의 주 요 농업형질의 상관은 Table 8과 같다. 이 표에서 보는 바와 같이 교잡종 자체의 수량구성의 모든 요소뿐만 아니라 간장, 착수고도 유의성을 보였고, 평균에 대한 잡종강세와는 종실수 량이 정의 상관을 보였다. 이것으로 옥수수 수량과 관련된 잡 종강세 예측이 가능할 것으로 사료되어 교배조합 구성을 위 한 노력 절감과 광대한 포장시험에 의한 시간과 노력을 어느 정도 감소시킬 수 있을 것으로 생각된다.
Table 8.
Correlation coefficients between genetic distance and F1, mid-parent heterosis (MPH) in 36 F1 hybrids based on selected 9 SSR primers and morphological traits.
Table 8.
Trait Correlation coefficients
F1 MPH
DA -0.14 -0.48**
DS -0.21 -0.37*
PH 0.60** -0.21
EH 0.61** -0.24
EL 0.46** 0.12
EW 0.34* 0.03
HW 0.33* -0.24
GY 0.70** 0.43**
반이면 교잡을 통해 얻어진 36개의 F1 교잡종들에 대하여 수량 및 농업관련 형질에 대하여 조사하였다. 36개 교배조합 의 F1과 교배친의 잡종강세를 조사한 결과, F1의 잡종강세 정 도는 종실수량에서 156%로 가장 컸고, 100립중이 가장 작은 7%로 나타났다. 농업형질에 의한 교배친들 사이의 표현형적 거리는 육성된 계보와 차이가 있었으며, 농업형질만으로 유전 적 거리를 측정하는 것은 한계가 있었다. SSR 분석에 의한 유전적 거리와 농업형질과의 상관을 확인하기 위하여 9개의 자식계통에 대한 92개의 SSR primer를 확인한 결과, 육성 계 보에 의한 유연관계를 명확하게 나타내지 못하여 표현형적 거리와 SSR 마커에 의한 유전적 거리에서도 유의성을 찾을 수 없었다. 본 연구에서는 옥수수 종실수량과 SSR 분석을 통 한 유전적 거리 사이의 유의성이 없었다. 따라서 종실수량과 정의 상관을 보이는 9개의 primer 조합을 선발하였다. 9개의 SSR primer들을 이용한 계통유연관계를 분석에서 옥수수 계 통들의 계보와 선발된 SSR을 이용한 유전적 거리는 연관성 을 찾을 수 없었지만, 종실수량과 선발된 9개의 SSR primer 를 이용한 유전적 거리와는 고도의 정의 상관을 보였다 (R2=0.703**). 선발된 SSR primer를 이용한 유전적 거리와 교잡종의 주요 농업형질의 상관 분석 결과, 교잡종 자체의 수 량구성의 모든 요소뿐만 아니라 간장, 착수고도 유의성을 보 였고, 평균에 대한 잡종강세와는 종실수량이 정의 상관을 보 였다. 본 연구에서 선발한 9개의 SSR primer들은 옥수수 육 종연구에서 교배조합 구성을 위한 잡종강세 예측에 도움을 줄 것으로 생각된다.
본 논문은 골든씨드프로젝트(식량종자사업단, 213001-04-3- SBA10, PJ009940022015)와 2015년도 강원대학교 대학회계 학술연구조성비(관리번호-520150106)의 지원에 의하여 연구 되었음.
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Genetic Analyses for Yield Components and Heterosis in Maize Elite Inbred Lines
Korean. J. Breed. Sci.. 2015;47(4):345-356.   Published online December 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.4.345
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Genetic Analyses for Yield Components and Heterosis in Maize Elite Inbred Lines
Korean. J. Breed. Sci.. 2015;47(4):345-356.   Published online December 31, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.4.345
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Genetic Analyses for Yield Components and Heterosis in Maize Elite Inbred Lines
Image Image Image
Fig. 1. Correlations between mid-parent heterosis (MPH), average yields of parents and grain yields (GY) in F1 hybrids of 36 cross combinations.
Fig. 2. An example of an SSR profile of 9 maize inbred lines.
Fig. 3. Cluster diagram among the 9 maize inbred lines based on average phenetic distance (a), genetic distance of 92 SSR primers (b), genetic distance of 9 SSR primers (c).

Fig. 1.

Fig. 2.

Fig. 3.

Genetic Analyses for Yield Components and Heterosis in Maize Elite Inbred Lines

Maize elite inbred lines used in this study.

Class Line
Lancaster Sure Crop (LSC) Mo17, C103, Oh43, Va85
Iowa Stiff Stalk Synthetic (BSSS) B73, Wf9, B14A, A632
Others ND203

Morphological traits measured on individual maize plants.

Traits Abbr. Units Remarks
Days to Anthesis DA day Days from sowing to pollen shed
Days to Silking DS day Days from sowing to silk emergence
Plant Height PH cm Height from soil level to top-leaf node at maturity
Ear Height EH cm Height from soil level to top-ear node at maturity
H. turcicum Ht 0~9 0: very strong resistant, 9: very weak susceptible
Corn Borer CB 0~9 0: very strong resistant, 9: very weak susceptible
Stalk Rot SR 0~9 0: very strong resistant, 9: very weak susceptible
Stalk Lodging SL 1~9 1: no lodging, 9: totally lodging
Ear Length EL cm Length of ear
Ear Width EW cm Width of ear
100 kernel Weight HW g Weight of 100 kernels at 15.5% moisture content
Grain Yield GY kg/10a Grain yields per 10a at 15.5% moisture content

Means of 8 agronomic traits in 9 maize inbred lines.

Inbred line DA (day) DS (day) PH (cm) EH (cm) EL (cm) EW (cm) HW (g) GY (kg/10a)
A632 88 90 148 67 9.8 3.1 27.1 395
ND203 88 90 199 101 10.8 2.8 28.9 281
Mo17 86 90 153 66 13.8 3.4 32.2 457
Oh43 82 85 166 70 16.9 3.4 24.4 385
C103 82 85 141 56 13.2 3.1 29.1 423
B14A 86 91 159 69 12.2 3.4 27.1 246
Va85 86 89 209 105 16.2 3.8 39.7 389
Wf9 86 89 203 109 11.8 3.3 30.3 347
B73 88 90 159 65 11.2 3.6 33.4 334

Means of 12 agronomic traits in 36 cross combinations.

Cross Combination DA (day) DS (day) SL (1-9) Ht (0-9) SR (0-9) CB (0- PH (cm) EH (cm) EL (cm) EW (cm) HW (g) GY (kg/10a)
A632/ND203 73 76 2 4 3 3 226 112 16.3 4.1 32.4 887
A632/Mo17 73 76 1 0 0 0 247 130 17.7 4.2 31.5 955
A632/Oh43 72 76 1 4 4 4 229 106 15.8 3.9 31.5 845
A632/C103 73 76 1 0 0 0 226 105 16.2 4.3 26.2 808
A632/B14A 73 76 1 6 4 4 233 111 14.8 4.1 32.6 721
A632/Va85 74 76 3 5 4 4 242 124 18.3 4.5 32.1 1,015
A632/Wf9 73 76 1 6 4 4 247 125 16.9 4.5 31.9 977
A632/B73 75 78 2 7 4 4 228 107 15.1 4.4 30.9 844
ND203/Mo17 73 76 1 0 0 0 239 132 17.6 4.2 31.1 849
ND203/Oh43 73 76 1 0 0 0 246 127 18.9 4.3 33.4 824
ND203/C103 73 76 1 0 0 0 233 117 18.6 4.2 32.3 859
ND203/B14A 73 77 4 5 4 5 251 134 18.1 4.4 34.2 944
ND203/Va85 77 79 1 0 0 0 241 122 17.5 4.4 31.1 968
ND203/Wf9 73 76 6 6 4 4 255 136 16.4 4.4 33.9 862
ND203/B73 72 76 1 2 2 1 243 128 17.0 4.3 33.0 888
Mo17/Oh43 72 75 1 4 4 4 232 109 18.1 4.3 34.3 889
Mo17/C103 73 76 1 0 0 0 229 106 17.5 4.1 27.8 802
Mo17/B14A 73 76 1 5 4 4 248 122 18.1 4.6 36.4 1,097
Mo17/Va85 73 76 3 4 4 5 247 119 19.7 4.5 34.3 1,080
Mo17/Wf9 72 75 2 6 4 4 263 133 17.6 4.5 35.0 1,068
Mo17/B73 74 77 2 6 4 5 225 104 18.2 4.3 33.4 805
Oh43/C103 74 77 1 2 1 2 224 108 17.1 4.1 29.1 816
Oh43/B14A 74 77 1 5 4 4 219 103 16.6 4.0 31.3 757
Oh43/Va85 72 75 1 4 4 4 244 115 20.0 4.4 36.1 995
Oh43/Wf9 75 76 1 0 0 0 259 134 16.5 4.2 28.7 864
Oh43/B73 73 76 1 5 4 4 237 112 18.7 4.3 34.4 876
C103/B14A 74 76 1 3 3 3 238 110 17.5 4.4 29.3 896
C103/Va85 73 76 3 5 4 4 245 118 18.2 4.4 29.4 889
C103/Wf9 73 76 2 7 4 4 266 134 16.8 4.6 31.1 975
C103/B73 74 77 1 5 4 5 236 108 17.5 4.4 29.4 759
B14A/Va85 74 77 3 4 3 4 255 120 18.7 4.6 34.4 1,045
B14A/Wf9 73 77 2 3 3 2 258 139 17.8 4.5 35.5 978
B14A/B73 75 77 1 6 4 5 228 103 17.1 4.3 31.2 938
Va85/Wf9 73 75 5 4 3 3 269 143 18.4 4.5 31.0 1,017
Va85/B73 74 76 5 4 4 4 240 112 19.1 4.4 32.8 1,064
Wf9/B73 73 76 1 8 4 3 272 142 16.3 4.6 32.5 1,012

Means and ranges of F1 hybrids and their mid-parent heterosis (MPH) in 36 cross combinations.

Trait F1 MPH (%)
Means Ranges Means Ranges
DA 73 72~77 -14 -17~-10
DS 76 75~79 -14 -17~-9
PH 242 219~272 43 18~64
EH 120 103~143 54 18~95
EL 17.5 14.8~20.0 26 7~44
EW 4.4 3.9~5.5 21 10~52
HW 32.1 26.2~36.4 7 -15~25
GY 913 721~1,097 156 82~258

※MPH (Mid-Parent Heterosis) = (F1-((P1+P2)/2))/(P1+P2)/2)×100

GCA and SCA effects of grain yields calculated in half-diallel cross of 9 maize inbred lines.

SCA effects GCA effects
ND203 Mo17 Oh43 C103 B14A Va85 Wf9 B73
A632 41.9 43.6 30.6 2.4 -166.3 28.1 35.9 -16.1 -36
ND203 -66.6 5.4 49.3 52.6 -23 -83.3 23.7 -31.9
Mo17 4.1 -74 139.3 22.7 56.4 -125.6 34.4
Oh43 37 -103.7 34.7 -50.6 42.4 -62.6
C103 44.1 -62.4 69.3 -65.7 -71.4
B14A 11.9 -9.4 31.6 10.3
Va85 -70 58 109.9
Wf9 51.7 64.1
B73 -16.9

Number of allelic bands and PIC values of the SSR loci among 9 maize inbred lines.

Locus No. of allele PICz Locus No. of allele PIC Locus No. of allele PIC Locus No. of allele PIC
phi014 2 0.2 bnlg589 4 0.67 phi053 2 0.8 phi075 2 0.21
phi016 2 0.49 bnlg439 6 0.75 phi057 3 0.37 phi081 4 0.2
phi027 2 0.49 phi011 2 0.64 phi059 2 0.44 phi091 5 0.33
phi031 3 0.53 phi019 6 0.75 phi076 6 0.27 phi092 4 0.38
phi037 3 0.73 phi006 7 0.79 phi077 7 0.59 phi102 4 0.51
phi047 3 0.68 phi037 3 0.73 phi078 4 0.2 phi040 2 0.51
phi073 2 0.35 phi086 4 0.22 phi079 3 0.77 phi058 3 0.16
phi083 2 0.49 phi021 5 0.8 phi080 4 0.56 phi061 4 0.36
phi087 2 0.49 phi026 5 0.77 phi082 3 0.53 phi062 4 0.25
phi090 2 0.51 phi032 2 0.49 phi084 2 0.69 phi085 6 0.85
phi119 4 0.72 phi098 4 0.49 phi096 3 0.09 phi093 3 0.09
phi127 3 0.57 phi113 6 0.7 phi097 2 0.44 phi117 3 0.38
bnlg278 4 0.62 phi125 3 0.64 phi028 5 0.81 phi118 6 0.35
bnlg386 2 0.78 phi033 3 0.57 phi039 4 0.81 phi049 2 0.1
bnlg161 6 0.8 phi034 3 0.57 phi112 4 0.52 phi116 3 0.44
bnlg108 4 0.73 phi035 2 0.44 phi114 2 0.21 phi121 4 0.49
bnlg125 6 0.64 phi038 2 0.49 phi054 5 0.79 phi123 6 0.56
bnlg166 5 0.69 phi041 3 0.57 phi056 7 0.69 phi068 5 0.8
bnlg420 6 0.51 phi044 2 0.35 phi063 3 0.57 phi126 5 0.8
bnlg210 5 0.77 phi045 2 0.78 phi064 3 0.68 phi129 4 0.6
bnlg594 4 0.84 phi089 6 0.7 phi065 7 0.51 2-7 0.09-0.85
bnlg640 4 0.81 phi048 3 0.53 phi066 2 0.44 Means 3.71 0.55
bnlg155 5 0.8 phi050 3 0.73 phi067 4 0.36
bnlg398 2 0.54 phi051 4 0.64 phi072 2 0.69

zPIC : Polymorphic Information Content.

Correlation coefficients between genetic distance and F1, mid-parent heterosis (MPH) in 36 F1 hybrids based on selected 9 SSR primers and morphological traits.

Trait Correlation coefficients
F1 MPH
DA -0.14 -0.48**
DS -0.21 -0.37*
PH 0.60** -0.21
EH 0.61** -0.24
EL 0.46** 0.12
EW 0.34* 0.03
HW 0.33* -0.24
GY 0.70** 0.43**
Table 1. Maize elite inbred lines used in this study.
Table 2. Morphological traits measured on individual maize plants.
Table 3. Means of 8 agronomic traits in 9 maize inbred lines.
Table 4. Means of 12 agronomic traits in 36 cross combinations.
Table 5. Means and ranges of F1 hybrids and their mid-parent heterosis (MPH) in 36 cross combinations.

MPH (Mid-Parent Heterosis) = (F1-((P1+P2)/2))/(P1+P2)/2)×100

Table 6. GCA and SCA effects of grain yields calculated in half-diallel cross of 9 maize inbred lines.
Table 7. Number of allelic bands and PIC values of the SSR loci among 9 maize inbred lines.

PIC : Polymorphic Information Content.

Table 8. Correlation coefficients between genetic distance and F1, mid-parent heterosis (MPH) in 36 F1 hybrids based on selected 9 SSR primers and morphological traits.

Table 1.

Table 2.

Table 3.

Table 4.

Table 5.

Table 6.

Table 7.

Table 8.

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