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듀럼밀 유전자원의 질소시비 수준에 따른 농업형질 변이 분석

천호선1, 곽순화1, 최시은1, 이수경2, 박진희3, 김경민3, 박철수1, 모영준1,4,*

Variations in Major Agronomic Traits of Durum Wheat Germplasm Under Different Nitrogen Fertilization Levels

Hosun Cheon1, Sun-Hwa Kwak1, Sieun Choi1, Sukyeung Lee2, Jinhee Park3, Kyung-Min Kim3, Chul Soo Park1, Youngjun Mo1,4,*
Korean Journal of Breeding Science 2024;56(3):281-291.
Published online: September 1, 2024

1전북대학교 농업생명과학대학 작물생명과학과

2농촌진흥청 기술협력국 국제기술협력과

3농촌진흥청 국립식량과학원

4전북대학교 농업과학기술연구소

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*Corresponding to Youngjun Mo TEL. +82-63-270-2530 E-mail. yjmo@jbnu.edu
• Received: July 9, 2024   • Revised: August 6, 2024   • Accepted: August 6, 2024

Copyright © 2024 by the Korean Society of Breeding Science

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  • Breeding wheat cultivars with high nitrogen use efficiency is crucial for sustainable agriculture. In this study, 130 durum wheat accessions from 53 countries were cultivated under two nitrogen fertilization levels to investigate variations in agronomic traits and identify accessions suitable for low-nitrogen conditions. Under no nitrogen fertilization, SPAD value (-99.2), flag leaf length (−15.8 mm), grain area (−0.52 mm2), grain length (−0.27 mm), thousand-grain weight (+5.2 g), and grain protein content (−4.2%) showed significant differences compared to that under standard nitrogen fertilization. Among these traits, the SPAD value exhibited a strong positive correlation with protein content in the absence of nitrogen fertilization, suggesting its potential as an indicator for selecting germplasm with superior nitrogen use efficiency. Hierarchical cluster analysis, based on the differences in the six traits under the two nitrogen fertilization levels, classified the 130 durum wheat accessions into three groups (Groups 1, 2, and 3). Accessions in Group 2 exhibited superior adaptability under no nitrogen fertilization compared with those in the other two groups. We identified seven accessions in Group 2 that exhibited minimal decreases in SPAD values and protein content under no nitrogen fertilization for use in wheat breeding programs aimed at developing cultivars with high nitrogen use efficiencies.
밀은 인류의 영양 공급에 높은 비중을 차지하는 가장 중요한 곡물 중 하나이며(Colasuonno et al. 2019, Hong et al. 2023) 그 중 4배체(AABB) 듀럼밀(Triticum turgidum spp. durum)은 세계 밀 생산의 5%를 차지한다(Beres et al. 2020). 듀럼밀은 6배체 보통밀(Triticum aestivum, common wheat, AABBDD)보다 종실이 단단하고 높은 단백질 함량을 나타내며 주로 파스타 제조에 사용되어 지중해 지역에서 많이 재배되고 있다(Mastrangelo & Cattivelli 2021).
듀럼밀의 품질에는 단백질 함량이 중요하기 때문에 재배 기간 중 질소 시비가 필수적이다. 질소는 단백질 합성, 기관 발달 등 작물 생장에 필수적인 영양소로 질소 시비는 작물에서 여러 가지 농업형질에 관여하여 수량에 큰 영향을 준다. 일반적으로 질소 시비는 식물체 잎의 엽록소 함량과 엽면적 지수를 증가시킴으로써 광합성량을 증가시키고 종실 크기와 전분, 단백질 함량 등에 관여한다(Amanullah et al. 2009, Cui et al. 2023, Heo & Sherman 2012, Lyngstad 1975, Mălinaş et al. 2022). 밀은 시비되는 질소의 30~40% 정도만 흡수하며 나머지는 대기나 토양으로 유출되는 것으로 추정된다(Lyngstad 1975, Nguyen et al. 2016). 밀의 수요 증가에 따라 질소 비료 사용량도 지속적으로 증가해왔으며, 과도한 질소 시비는 온실가스 발생, 토양 및 지하수로의 질소 유출 등 심각한 환경 오염을 유발할 수 있다(Tilman 1999). 농업에서 질소 비료의 온실가스 배출 관여도는 75%에 육박하며 질산염 유출로 인한 토양 오염의 문제도 심각해지고 있다(Mălinaş et al. 2022, Salim & Raza 2020). 따라서 늘어나는 밀 수요를 충족함과 동시에 질소 시비량을 최소화하여 지속 가능한 농업을 실현하는 것이 중요하다.
듀럼밀은 일반적으로 보통밀보다 생체중과 종자 단백질 함량이 높으며, 유전자형에 따라 질소이용효율에 차이가 있으므로 듀럼밀 유전자원 중 저질소 조건에서 효과적으로 질소를 흡수하고 이용할 수 있는 유전자형을 식별하는 것은 질소 비료 사용 감소를 위한 육종에서 중요하다(Gaju et al. 2011, Liang et al. 2014). 본 연구는 50여 개국 유래 130개 듀럼밀 유전자원을 질소 비료 무시비구와 표준시비구 2가지 조건에서 재배하여 다양한 농업형질 변이를 조사하고 형질별로 질소 수준에 영향을 받는 정도를 평가하였으며, 추후 질소이용효율 증대 품종개발에 활용될 수 있는 후보 유전자원들을 선발하였다.
시험재료 및 재배조건
농촌진흥청 농업유전자원센터로부터 분양 받은 러시아, 에티오피아, 불가리아 등 53개국 유래 130개 듀럼밀 유전자원을 연구에 사용하였다. 시험재료는 전북대학교 시험포장에 2022년 10월 21일에 파종하였으며, 휴립조파로 자원당 종자 25립을 1열(1.5 m)에 자원 간 30 cm 간격으로 파종하였다. 표준시비구는 농촌진흥청 표준재배법(RDA 2020)을 기준으로 질소를 기비 3.6 kg/10a, 수비 5.5 kg/10a로 분시하였고, P2O5를 7.4 kg/10a, K2O를 3.9 kg/10a 기비로 시비하였다. 무시비구에는 질소 비료를 시비하지 않았으며 표준시비구와 같은 양의 P2O5, K2O만을 기비로 시비하였다. 기타 재배관리는 농촌진흥청 표준 재배법에 준하여 실시하였으며, 표준 품종으로 금강을 매 20열마다 1열씩 재식하여 농업형질을 비교하였다.
농업형질조사
출수일수(days to heading, DH)는 파종일부터 총 경수의 40% 이상이 출수한 날까지의 일수를 조사하였다. 출수 후 25일에 지엽 중간 부분에서 SPAD값(SPAD-502, Minolta Ltd, Osaka Japan), 지엽길이(flag leaf length, FL), 지엽 폭(flag leaf width, FW)을 5반복 측정하였다. 1수소수수(spikelet number per spike, SNS)는 6-7월 성숙기에 자원 당 5수로부터 측정하였고, 같은 시기에 간장(culm length, CL), 수장(spike length, SL), 망장(awn length, AL)도 조사하였다. 종자 수확 이후 ImageJ 프로그램을 사용하여 종실면적(grain area, GA), 종실길이(grain length, GL), 종실폭(grain width, GW)을 100립씩 2반복 측정하였으며, 천립중(thousand grain weight, TGW)은 200립의 무게를 측정하고 1,000립 무게로 환산하여 계산하였다. 종실 단백질함량(protein content, PC)은 Elementar rapid MAX N exceed를 사용하여 분쇄한 종실의 단백질 함량을 자원 당 3반복 측정하였다.
통계분석
통계분석은 R (Version 4.0.2, The R Foundation for Statistical Computing Platform)을 이용하였으며, 두 시비구 간 유전자원의 농업형질 평균 등의 기술통계와 t-test를 이용한 변이 분석은 ‘stats’ 패키지를 사용하였다. 농업형질 간의 상관분석은 ‘corrplot’ 패키지, 두 시비구간 농업형질 분포 확인은 ‘ggplot2’ 패키지, 자원 간 PCA 분석을 통한 다변량 변수의 분석은 ‘ggfortigy’ 패키지를 사용하였다. UPGMA Hierarchical Clustering은 ‘ggplot2’ 패키지를 사용하였고, 형질 간 상관관계를 이용하여 130 자원을 그룹화한 덴드로그램을 시각화하였다. ANOVA와 그룹 간 평균 비교를 위한 Tukey 분석은 ‘agricolae’ 패키지를 이용하였다.
질소 시비에 따른 농업형질 변이 분석
듀럼밀 130 유전자원의 표준시비구 및 무시비구에서 출수일수, 간장, 수장, 망장, SPAD값, 지엽길이, 지엽폭, 1수소수수, 종실면적, 종실길이, 종실폭, 천립중, 단백질함량을 비교한 결과는 Table 1과 같다. 표준시비구와 무시비구 간 t-test 결과 SPAD값, 지엽길이, 종실면적, 종실길이, 천립중, 단백질함량은 유의한 차이를 보였으며(p<0.05), 출수일수, 간장, 수장, 망장, 1수소수수, 지엽폭, 종실폭은 유의한 차이가 없었다. 평균 SPAD값은 표준시비구에서 552.2 (451.0-679.0), 무시비구에서 453.0 (303.2-569.0)으로 18.0% 감소하였다. 지엽길이는 표준시비구 265.4 mm (196.0-390.6 mm), 무시비구 249.6 mm (135.0-346.2 mm)로 6.0% 감소하였다. 종실면적은 표준시비구 15.73 mm2 (11.37-21.56 mm2), 무시비구 15.21 mm2 (10.82-21.51 mm2)로 3.3% 감소하였다. 종실길이는 표준시비구 6.87 mm (5.60-9.28 mm), 무시비구 6.60 mm (5.46-8.04 mm)로 4.1% 감소하였다. 한편 천립중은 표준시비구 39.3 g (20.7-60.8 g), 무시비구 44.5 g (26.9- 74.3 g)으로 무시비구에서 13.4% 무거웠다. 종실 단백질함량은 두 시비구 간에 가장 큰 차이를 보였는데(p<0.0001***) 표준시비구 평균 17.0% (13.5-22.7%), 무시비구 평균 12.8% (9.5-19.2%)으로 무시비구에서 25.6% 감소하였다. 대조구로 사용된 금강은 표준시비구 대비 무시비구에서 SPAD값 29.1% (531.0→376.4), 종실면적 2.8% (19.19→18.65 mm2), 종실길이 3.7% (7.21→6.94 mm), 단백질함량 33.9% (18.0→11.9%)가 감소하였다.
질소 시비에 따른 농업형질 상관분석
두 시비조건에서 각각 13가지 농업형질들 간의 상관분석을 수행하였으며(Fig. 1), |r|≥0.30의 유의한 상관관계를 나타낸 형질들은 다음과 같다. 두 시비구에서 모두 간장은 1수소수수와 양의 상관(r=0.38-0.40)을 나타냈으며, 망장도 지엽길이, 종실면적, 종실길이, 천립중과 양의 상관(r=0.37-0.48)을 나타냈다. 지엽폭은 SPAD값(r=0.31-0.42)과 지엽길이(r=0.40-0.49)와 양의 상관을 보였으며, 그 중 지엽길이와 지엽폭은 종실면적과도 양의 상관(r=0.34-0.42)을 보였다. 종실면적은 종실길이, 종실폭과 강한 양의 상관(r=0.75-0.88)을 보였으며 종실길이와 종실폭은 상대적으로 약한 상관(r=0.31-0.40)을 보였다. 종실면적, 종실길이, 종실폭은 모두 천립중과 강한 양의 상관(r=0.61-0.87)을 보였다. 반면 시비구 간 상관관계의 양상이 다른 형질들도 있었는데, 간장은 표준시비구에서만 종실면적(r=0.34), 종실폭(r=0.41), 천립중(r=0.36)과 양의 상관을 보였다. SPAD값은 무시비구에서 지엽길이(r=0.30), 단백질 함량(r=0.50)과 양의 상관을 나타냈지만 표준시비구에서는 유의한 상관관계가 나타나지 않았다.
각 형질에 대하여 유전자원별로 표준시비구 대비 무시비구에서의 변화량(무시비구 값 - 표준시비구 값)을 수치화하여 상관분석을 수행한 결과는 Fig. 2와 같다. 출수일수 변화량은 간장 변화량(r=-0.36) 및 SPAD 값 변화량(r=-0.36)과 음의 상관을 보였으며, 천립중 변화량과 단백질함량 변화량 사이에도 음의 상관(r=-0.41)이 나타났다. 반면 SPAD값 변화량은 지엽길이(r=0.38), 단백질함량(r=0.45)의 변화량과 양의 상관을 나타냈다. 또한 지엽폭의 변화량은 지엽길이(r=0.68), 1수소수수(r=0.41), 종실길이(r=0.31)의 변화량과 양의 상관을 나타냈고, 종실면적의 변화량은 종실길이(r=0.68), 종실폭(r=0.83), 천립중(r=0.54)의 변화량과 강한 양의 상관을 보였으며, 천립중과 종실폭의 변화량도 강한 양의 상관을 보였다(r=0.53).
시비구 간 농업형질 변화에 따른 유전자원의 군집분석
표준시비구와 무시비구 간 t-test에서 유의한 차이를 보인 6개 형질(SPAD값, 지엽길이, 종실면적, 종실길이, 천립중, 단백질함량)의 변화량을 기준으로 계층적 군집분석을 수행한 결과, 130개 듀럼밀 유전자원은 Group 1 (n=2), Group 2 (n=87), Group3 (n=41) 3개의 그룹으로 분류할 수 있었다(Fig. 3a). 주성분분석(principal component analysis; PCA) 결과 PC1 (36.2%)과 PC2 (30.4%)는 전체 변이의 66.6%를 설명하였으며(Fig. 3b) PC1을 기준으로 모든 형질들은 양의 방향성을, PC2에 대해서는 SPAD값, 지엽길이, 단백질함량이 양의 방향성을, 종실면적, 종실길이, 천립중은 음의 방향성을 나타냈다. 군집분석에 따른 세 그룹 간 ANOVA를 통해 6개 형질의 평균을 비교하였을 때 표준시비구에선 그룹 간 유의한 차이가 나타나지 않았다(Table 2). 반면 무시비구에서는 SPAD값과 단백질함량이 그룹 간 유의한(p<0.0001) 차이를 보였고, 두 형질 모두 Group 1, 2, 3 순으로 높은 값을 보였다.
표준시비구 대비 무시비구에서의 형질 변화량은 SPAD값이 3개 그룹 간에 유의한 차이를 나타냈으며, Group 1, 2, 3 순으로 저질소 조건에서 낮은 감소를 보였다. 단백질 함량과 종실길이는 Group 2, 3 간 유의한 차이를 나타냈으며, Group 2가 3보다 낮은 감소를 보였다. 지엽길이는 Group 1, 2와 Group 3 간 유의한 차이를 나타냈으며, Group 1, 2가 3보다 낮은 감소를 보였다(Table 2, Fig. 4).
본 연구에서는 130개 듀럼밀 유전자원을 서로 다른 질소 시비 조건에서 재배하고 주요 농업 형질의 변이를 분석하였다. 밀의 질소 이용 능력은 이전의 여러 연구에서 유전자형에 따라 차이가 존재한다는 것이 보고되었다(Gaju et al. 2011, Liang et al. 2014). 본 연구에서 두 시비구 간 유의한 차이를 나타낸 형질 6개(SPAD값, 지엽길이, 종실넓이, 종실길이, 천립중, 종실 단백질 함량) 중 특히 큰 차이를 보였던 것은 SPAD값, 천립중, 종실 단백질 함량이었다(p<0.0001***, Table 1). SPAD값은 식물체 엽록소 함량을 측정함으로써 식물체의 N 수준을 비파괴적인 방법으로 예측할 수 있는 유용한 지표이다. 일반적으로 식물체의 N 함량을 측정하기 위해서는 전체 식물체를 채집해야 하지만 SPAD를 사용한다면 지엽의 SPAD 측정만으로 식물체의 N 수준을 예측할 수 있다(Asplund et al. 2016, Monostori et al. 2016, Xiong et al. 2015). 밀의 지엽은 이삭 바로 아래의 잎으로써 종자 N 함량과 천립중에 크게 관여하므로(Khaliq et al. 2008), 지엽에서 높은 SPAD값과 큰 지엽면적을 나타내는 밀 품종은 종실 단백질 함량 또한 높게 나타나는 경향을 보인다(Monostori et al. 2016).
본 연구의 무시비구에서 종자 단백질 함량은 SPAD값, 지엽길이와 양의 상관을 나타냈으나 표준시비구에서는 이러한 상관관계가 나타나지 않았는데(Fig. 1), 이는 최적 질소조건에서는 품종 간 SPAD값의 차이에 따른 종실단백질 함량 변이가 나타나지 않는다는 것을 시사한다. 듀럼밀 종자의 단백질 함량은 평균 12%-16% 정도이며(Saini et al. 2023), 식물 종자 내부에서 전분 등과 함께 일정 비율로 존재한다. 토양 질소가 충분한 조건에서는 식물체의 질소 함량이 최적의 종실 단백질 함량을 채우는데 충분했기 때문에 이런 결과가 나타난 것으로 추측된다. 실제로 Monostori et al. (2016)의 연구에서는 서로 다른 질소 조건에서 SPAD값을 통해 밀의 수량관련 지표들을 예측했을 때 저질소 조건에서만 이러한 예측이 가능한 것을 확인하였다. 본 연구에서도 무시비구에서 SPAD값과 종실 단백질 함량은 양의 상관을 나타냈기 때문에 무시비구의 SPAD값은 질소이용효율이 높은 유전자원을 선발하는 지표로 이용할 수 있을 것으로 판단된다.
일반적으로 식물이 질소 결핍 스트레스를 받게 되면 식물체 생체중이 감소하고 수량구성요소 또한 감소하는 경향을 보인다(Wang et al. 2021). 본 연구에서 천립중은 무시비구에서 표준시비구보다 오히려 더 높은 경향을 보였는데, 저질소 조건에서 천립중 증가는 스트레스 상황에서 종실 내 세포벽, 셀룰로오스 축적 증가로 인한 종실 크기 증가의 결과임이 보고된 바 있다(Wang et al. 2021). 또한 저질소 조건에서는 단위면적 당 이삭 수 감소로 인하여 이에 대한 상쇄효과(trade-off)로 천립중 증가가 발생한 가능성도 있는 것으로 고찰된다(Kim & Kim 2023). 본 연구에서 수수 측정은 이루어지지 않았으므로, 추후 연차반복 실험에서는 수수 및 1수립수의 측정도 병행하여 질소시비에 따른 수량구성요소의 변화를 보다 면밀하게 살펴볼 계획이다.
본 연구에서는 130개 듀럼밀 유전자원 중 저질소 조건에서 적응성이 우수하다고 판단되는 7개 후보 유전자원을 선발하였다(Table 3). 선발 자원들은 무시비구에서 출수일수가 200일 이하이며 간장이 100 cm 이하로 국내 환경 조건에서 최소한의 적응성을 보유하였다고 판단되는 자원들 중, 표준시비구 대비 무시비구에서 SPAD값과 단백질 함량의 감소가 최소인 자원들을 선정하였다. 선정한 자원들은 모두 SPAD값과 단백질 함량의 변화율이 가장 작게 나타난 Group 2에 속해 있었으며(Table 2), 표준시비구 대비 무시비구에서의 SPAD값 감소는 평균 -43.5, 단백질 함량 감소는 평균 -2.7%로 대조구 품종 금강이 각각 -154.6, -6.1% 감소한 것과 비교하여 감소량이 현저히 작았다(Table 3). 해당 유전자원들은 2년차 연차 반복 조사를 통해 환경변이를 조사할 예정이며 주당수수, 일수립수 등 다른 수량 구성요소들에 대한 추가 조사를 수행하여 질소조건에 따른 수량변화를 보다 면밀하게 검정할 예정이다. 또한 추후 표현형 데이터와 유전자형 데이터를 이용한 GWAS 분석을 통해 질소 이용 능력과 관련된 연관 유전자위를 탐색하여 질소 비료 저감을 위한 밀 육종사업에 활용하고자 한다.
본 연구는 저질소 조건에서 적응성이 우수한 듀럼밀 유전자원 선발을 위하여 50여개국 유래 130개 듀럼밀 유전자원을 질소 비료 무시비구와 표준시비구 조건에서 재배하고 13가지 농업형질(출수일수, 간장, 수장, 망장, SPAD값, 지엽길이, 지엽폭, 1수소수수, 종실면적, 종실길이, 종실폭, 천립중, 단백질함량)의 변이를 조사하였다. 표준시비구 대비 무시비구에 유의한 차이를 보인 형질은 SPAD값(-99.2), 지엽길이(-15.8 mm), 종실면적(-0.52 mm2), 종실길이(-0.27 mm), 천립중(+5.2 g), 단백질함량(-4.2%)으로, 특히 SPAD값은 무시비구에서의 종실 단백질함량과 강한 상관관계를 보여 질소이용능력 우수 자원 선발을 위하여 유용한 지표로 이용할 수 있음을 확인하였다. 두 시비구간 유의한 차이를 보인 6가지 형질의 변화량을 이용하여 군집분석을 수행한 결과 130개 유전자원을 3개 그룹(Group 1, 2, 3)으로 분류할 수 있었고, Group 2에 속한 자원들이 다른 두 그룹에 비하여 무시비구에서의 적응성이 우수한 것으로 평가되었다. 이 중 무시비구에서 출수일수가 200일 이하이며 간장이 100 cm 이하로 국내 환경 조건에서 최소한의 적응성을 보유하면서 무시비구에서 SPAD값과 종실 단백질함량 감소가 최소인 유전자원 7개를 선발하였으며, 향후 이들을 질소비료 저감을 위한 밀 육종사업에 활용하고자 한다.
본 연구는 농촌진흥청 공동연구사업(RS-2024-00432490)의 지원으로 수행되었습니다.
Fig. 1
Correlation analysis of 13 agronomic traits under different nitrogen fertilizations. a) standard nitrogen fertilization, b) no nitrogen fertilization. Each trait’s distribution is depicted along the diagonal, and beneath and above it, there are scatter plots featuring a fitted line and correlation coefficients (r), respectively. DH : days to heading, CL : culm length, SL : spike length, AL : awn length, SPAD : SPAD value, FL : flag leaf length, FW : flag leaf width, SNS : spikelet number per spike, GA : grain area, GL : grain length, GW : grain width, TGW : thousand grain weight, PC : protein content.
kjbs-56-3-281-f1.jpg
Fig. 2
Correlation analysis of the difference of 13 agronomic traits between nitrogen fertilization levels. Each trait’s distribution is depicted along the diagonal, and beneath and above it, there are scatter plots featuring a fitted line and correlation coefficients (r), respectively. DH : days to heading, CL : culm length, SL : Spike length, AL : awn length, SPAD : SPAD value, FL : flag leaf length, FW : flag leaf width, SNS : spikelet number per spike, GA : grain area, GL : grain length, GW : grain width, TGW : thousand grain weight, PC : protein content.
kjbs-56-3-281-f2.jpg
Fig. 3
a) Hierarchical cluster analysis of 130 durum wheat accessions according to difference values of 6 agronomic traits in the no nitrogen fertilization relative to the standard nitrogen fertilization. Durum germplasms are classified by the hierarchical clustering (n=3) using ‘UPGMA’ method. b) Principal component analysis (PCA) of 130 durum wheat accessions according to change amounts of 6 agronomic traits. SPAD : SPAD value, FL : flag leaf length GA : grain area, GL : grain length, TGW : thousand grain weight, PC : protein content.
kjbs-56-3-281-f3.jpg
Fig. 4
Box plot of difference values for 6 agronomic traits among the classified groups. SPAD : SPAD value, FL : flag leaf length GA : grain area, GL : grain length, TGW : thousand grain weight, PC : protein content. ns : not significantly different at p<0.05, *** : significantly different at p<0.0001, ** : significantly different at p<0.001. Group average values with the same letter are not significantly different at p<0.05. (ANOVA followed by Tukey’s multiple comparison)
kjbs-56-3-281-f4.jpg
Table 1
Descriptive statistics of 13 agronomic traits in 130 durum wheat accessions under different nitrogen fertilizations.
Table 1
Trait Fertilizer Avez SDy Minx Maxw CVv Control
(Geumgang)
Day to heading
(Days)		 SNu 199.1 5.98 185 216 2.9 162
NNt 199.0 ns 5.45 186 215 2.7 162 ns
Culm length
(cm)		 SN 118.3 23.45 54.8 171.0 19.8 68.0
NN 113.2 ns 24.37 54.0 158.8 21.5 67.0 ns
Spike length
(cm)		 SN 8.4 1.44 5.33 13.0 17.0 8.4
NN 8.4 ns 2.55 4.60 20.2 30.4 8.3 ns
Awn length
(cm)		 SN 11.2 2.15 5.0 18.4 19.2 5.5
NN 10.7 ns 2.52 0.0 18.0 23.7 5.0 ns
SPAD SN 552.2 38.03 451.0 679.0 6.9 531.0
NN 453.0*** 54.84 303.2 569.0 12.1 376.4***
Flag leaf length
(mm)		 SN 265.4 33.68 196.0 390.6 12.7 195.4
NN 249.6** 35.38 135.0 346.2 14.2 194.2 ns
Flag leaf width
(mm)		 SN 16.8 2.74 10.0 24.2 16.4 11.0
NN 16.4 ns 2.26 10.8 22.4 13.8 11.0 ns
Spikelet number per spike SN 21.7 2.47 13.20 28.40 11.4 14.9
NN 21.6 ns 2.58 13.50 30.20 11.9 15.2 ns
Grain area
(mm2)		 SN 15.73 1.856 11.37 21.56 11.8 19.19
NN 15.21** 1.700 10.82 21.51 11.2 18.65**
Grain length
(mm)		 SN 6.87 0.571 5.60 9.28 8.3 7.21
NN 6.60*** 0.496 5.46 8.04 7.5 6.94**
Grain width
(mm)		 SN 2.91 0.184 2.42 3.41 6.3 3.37
NN 2.92 ns 0.171 2.40 3.40 5.8 3.40 ns
Thousand grain weight (g) SN 39.3 8.59 20.7 60.8 21.9 41.9
NN 44.5*** 7.44 26.9 74.3 16.7 42.0 ns
Protein content
(%)		 SN 17.0 1.60 13.5 22.7 9.4 18.0
NN 12.8*** 2.06 9.5 19.2 16.2 11.9***

zAve : average, ySD : standard deviation, xMin : minimum, wMax : maximum, vCV : coefficient of variation, uSN : standard nitrogen fertilization, tNN : no nitrogen fertilization, ns : not significantly different at p<0.05, ** : significantly different at p<0.001, *** : significantly different at p<0.0001.

Table 2
Agronomics traits of durum wheat germplasm under different nitrogen fertilizations according to three classified groups.
Table 2
Fertilizer Group SPADz FLy (mm) GAx (mm²) GLw (mm) TGWv (g) PCu (%)
SNt 1 487.0 289.8 18.17 7.54 44.8 18.8
2 546.9 263.4 15.85 6.87 40.2 17.0
3 569.0 270.1 15.45 6.87 37.1 17.0
ns ns ns ns ns ns
NNs 1 558.5 a 312.4 17.35 7.18 52.7 14.5 ab
2 477.8 b 255.3 15.39 6.64 45.2 13.4 a
3 396.8 c 235.1 14.80 6.48 43.0 11.3 b
*** ns ns ns ns ***
Difr 1 71.5 a 22.6 a -0.82 -0.36 ab 7.9 -4.27 ab
2 -69.1 b -8.1 a -0.46 -0.23 a 5.0 -3.64 a
3 -172.1 c -35.0 b -0.64 -0.38 b 5.9 -5.68 b
*** ** ns ** ns ***

zSPAD : SPAD value, yFL : flag leaf length xGA : grain area, wGL : grain length, vTGW : thousand grain weight, uPC : protein content, tSN : standard nitrogen fertilization, sNN : no nitrogen fertilization, rDif : difference values of traits under different fertilization condition, ns : not significantly different at p<0.05, ** : significantly different at p<0.001, *** : significantly different at p<0.0001, Values followed by the same letters within the same fertilizer category are not significantly different at p<0.05. (ANOVA followed by Tukey’s multiple comparison)

Table 3
Top seven accessions with superior adaptability under no nitrogen fertilization condition.
Table 3
IT. Number Name Origin DHz CLy (cm) ΔSPADx ΔTGWw (g) ΔPCv (%) Group
Control Geumgang Korea 162 67.0 -154.6 1.0 -6.1 3
306433 Hourah Pakistan 193 97.2 -8.2 1.1 -1.2 2
333402 CDSS04B00717T-0TOPY-41Y-0M-3Y-0M-1Y-0B Uzbekistan 195 73.4 -58.6 1.3 -1.4 2
307534 Valencia Serbia 193 99.2 -89.4 2.2 -1.9 2
330451 Cltr 8164 Ukraine 195 99.8 -78.8 -0.5 -3.3 2
333401 CDSS04B00716T-0TOPY-42Y-0M-1Y-0M-4Y-0B USA 193 71.2 -23.4 6.6 -3.6 2
198274 U0001099 Bulgaria 197 61.0 -21.0 14.3 -3.8 2
305863 ELS 6404-154 Italy 193 99.2 -25.4 10.9 -3.8 2
uAve 194 85.8 -43.5 5.1 -2.7

zDH : days to heading, yCL : culm length, xΔSPAD : SPAD difference value, wΔTGW : thousand grain weight difference value. vΔPC : protein content difference value, uAve : average value of the seven accessions.

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Variations in Major Agronomic Traits of Durum Wheat Germplasm Under Different Nitrogen Fertilization Levels
Korean. J. Breed. Sci.. 2024;56(3):281-291.   Published online September 1, 2024
DOI: https://doi.org/10.9787/KJBS.2024.56.3.281
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Variations in Major Agronomic Traits of Durum Wheat Germplasm Under Different Nitrogen Fertilization Levels
Korean. J. Breed. Sci.. 2024;56(3):281-291.   Published online September 1, 2024
DOI: https://doi.org/10.9787/KJBS.2024.56.3.281
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Variations in Major Agronomic Traits of Durum Wheat Germplasm Under Different Nitrogen Fertilization Levels
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Fig. 1 Correlation analysis of 13 agronomic traits under different nitrogen fertilizations. a) standard nitrogen fertilization, b) no nitrogen fertilization. Each trait’s distribution is depicted along the diagonal, and beneath and above it, there are scatter plots featuring a fitted line and correlation coefficients (r), respectively. DH : days to heading, CL : culm length, SL : spike length, AL : awn length, SPAD : SPAD value, FL : flag leaf length, FW : flag leaf width, SNS : spikelet number per spike, GA : grain area, GL : grain length, GW : grain width, TGW : thousand grain weight, PC : protein content.
Fig. 2 Correlation analysis of the difference of 13 agronomic traits between nitrogen fertilization levels. Each trait’s distribution is depicted along the diagonal, and beneath and above it, there are scatter plots featuring a fitted line and correlation coefficients (r), respectively. DH : days to heading, CL : culm length, SL : Spike length, AL : awn length, SPAD : SPAD value, FL : flag leaf length, FW : flag leaf width, SNS : spikelet number per spike, GA : grain area, GL : grain length, GW : grain width, TGW : thousand grain weight, PC : protein content.
Fig. 3 a) Hierarchical cluster analysis of 130 durum wheat accessions according to difference values of 6 agronomic traits in the no nitrogen fertilization relative to the standard nitrogen fertilization. Durum germplasms are classified by the hierarchical clustering (n=3) using ‘UPGMA’ method. b) Principal component analysis (PCA) of 130 durum wheat accessions according to change amounts of 6 agronomic traits. SPAD : SPAD value, FL : flag leaf length GA : grain area, GL : grain length, TGW : thousand grain weight, PC : protein content.
Fig. 4 Box plot of difference values for 6 agronomic traits among the classified groups. SPAD : SPAD value, FL : flag leaf length GA : grain area, GL : grain length, TGW : thousand grain weight, PC : protein content. ns : not significantly different at p<0.05, *** : significantly different at p<0.0001, ** : significantly different at p<0.001. Group average values with the same letter are not significantly different at p<0.05. (ANOVA followed by Tukey’s multiple comparison)

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Variations in Major Agronomic Traits of Durum Wheat Germplasm Under Different Nitrogen Fertilization Levels

Descriptive statistics of 13 agronomic traits in 130 durum wheat accessions under different nitrogen fertilizations.

Trait Fertilizer Avez SDy Minx Maxw CVv Control
(Geumgang)
Day to heading
(Days)		 SNu 199.1 5.98 185 216 2.9 162
NNt 199.0 ns 5.45 186 215 2.7 162 ns
Culm length
(cm)		 SN 118.3 23.45 54.8 171.0 19.8 68.0
NN 113.2 ns 24.37 54.0 158.8 21.5 67.0 ns
Spike length
(cm)		 SN 8.4 1.44 5.33 13.0 17.0 8.4
NN 8.4 ns 2.55 4.60 20.2 30.4 8.3 ns
Awn length
(cm)		 SN 11.2 2.15 5.0 18.4 19.2 5.5
NN 10.7 ns 2.52 0.0 18.0 23.7 5.0 ns
SPAD SN 552.2 38.03 451.0 679.0 6.9 531.0
NN 453.0*** 54.84 303.2 569.0 12.1 376.4***
Flag leaf length
(mm)		 SN 265.4 33.68 196.0 390.6 12.7 195.4
NN 249.6** 35.38 135.0 346.2 14.2 194.2 ns
Flag leaf width
(mm)		 SN 16.8 2.74 10.0 24.2 16.4 11.0
NN 16.4 ns 2.26 10.8 22.4 13.8 11.0 ns
Spikelet number per spike SN 21.7 2.47 13.20 28.40 11.4 14.9
NN 21.6 ns 2.58 13.50 30.20 11.9 15.2 ns
Grain area
(mm2)		 SN 15.73 1.856 11.37 21.56 11.8 19.19
NN 15.21** 1.700 10.82 21.51 11.2 18.65**
Grain length
(mm)		 SN 6.87 0.571 5.60 9.28 8.3 7.21
NN 6.60*** 0.496 5.46 8.04 7.5 6.94**
Grain width
(mm)		 SN 2.91 0.184 2.42 3.41 6.3 3.37
NN 2.92 ns 0.171 2.40 3.40 5.8 3.40 ns
Thousand grain weight (g) SN 39.3 8.59 20.7 60.8 21.9 41.9
NN 44.5*** 7.44 26.9 74.3 16.7 42.0 ns
Protein content
(%)		 SN 17.0 1.60 13.5 22.7 9.4 18.0
NN 12.8*** 2.06 9.5 19.2 16.2 11.9***

Agronomics traits of durum wheat germplasm under different nitrogen fertilizations according to three classified groups.

Fertilizer Group SPADz FLy (mm) GAx (mm²) GLw (mm) TGWv (g) PCu (%)
SNt 1 487.0 289.8 18.17 7.54 44.8 18.8
2 546.9 263.4 15.85 6.87 40.2 17.0
3 569.0 270.1 15.45 6.87 37.1 17.0
ns ns ns ns ns ns
NNs 1 558.5 a 312.4 17.35 7.18 52.7 14.5 ab
2 477.8 b 255.3 15.39 6.64 45.2 13.4 a
3 396.8 c 235.1 14.80 6.48 43.0 11.3 b
*** ns ns ns ns ***
Difr 1 71.5 a 22.6 a -0.82 -0.36 ab 7.9 -4.27 ab
2 -69.1 b -8.1 a -0.46 -0.23 a 5.0 -3.64 a
3 -172.1 c -35.0 b -0.64 -0.38 b 5.9 -5.68 b
*** ** ns ** ns ***

Top seven accessions with superior adaptability under no nitrogen fertilization condition.

IT. Number Name Origin DHz CLy (cm) ΔSPADx ΔTGWw (g) ΔPCv (%) Group
Control Geumgang Korea 162 67.0 -154.6 1.0 -6.1 3
306433 Hourah Pakistan 193 97.2 -8.2 1.1 -1.2 2
333402 CDSS04B00717T-0TOPY-41Y-0M-3Y-0M-1Y-0B Uzbekistan 195 73.4 -58.6 1.3 -1.4 2
307534 Valencia Serbia 193 99.2 -89.4 2.2 -1.9 2
330451 Cltr 8164 Ukraine 195 99.8 -78.8 -0.5 -3.3 2
333401 CDSS04B00716T-0TOPY-42Y-0M-1Y-0M-4Y-0B USA 193 71.2 -23.4 6.6 -3.6 2
198274 U0001099 Bulgaria 197 61.0 -21.0 14.3 -3.8 2
305863 ELS 6404-154 Italy 193 99.2 -25.4 10.9 -3.8 2
uAve 194 85.8 -43.5 5.1 -2.7
Table 1 Descriptive statistics of 13 agronomic traits in 130 durum wheat accessions under different nitrogen fertilizations.

zAve : average, ySD : standard deviation, xMin : minimum, wMax : maximum, vCV : coefficient of variation, uSN : standard nitrogen fertilization, tNN : no nitrogen fertilization, ns : not significantly different at p<0.05, ** : significantly different at p<0.001, *** : significantly different at p<0.0001.

Table 2 Agronomics traits of durum wheat germplasm under different nitrogen fertilizations according to three classified groups.

zSPAD : SPAD value, yFL : flag leaf length xGA : grain area, wGL : grain length, vTGW : thousand grain weight, uPC : protein content, tSN : standard nitrogen fertilization, sNN : no nitrogen fertilization, rDif : difference values of traits under different fertilization condition, ns : not significantly different at p<0.05, ** : significantly different at p<0.001, *** : significantly different at p<0.0001, Values followed by the same letters within the same fertilizer category are not significantly different at p<0.05. (ANOVA followed by Tukey’s multiple comparison)

Table 3 Top seven accessions with superior adaptability under no nitrogen fertilization condition.

zDH : days to heading, yCL : culm length, xΔSPAD : SPAD difference value, wΔTGW : thousand grain weight difference value. vΔPC : protein content difference value, uAve : average value of the seven accessions.

Table 1

Table 2

Table 3

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