Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (5): 985-995.doi: 10.3864/j.issn.0578-1752.2026.05.005

• TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY·AGRICULTURE INFORMATION TECHNOLOGY • Previous Articles     Next Articles

Multi-Model Elucidating of Nutritional Quality Contributions to Maize Kernel Test Weight and Regional Heterogeneity

DONG JinLong1,2(), ZHAO Ying2, YU HaiBing1, LÜ JianYe2, QIN JiaQi2, LIANG Chen2, MING Bo2, LI ShaoKun1,2()   

  1. 1 Anhui Science and Technology University, Fengyang 233100, Anhui
    2 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences/State Key Laboratory of Crop Gene Resources and Breeding, Beijing 100081
  • Received:2025-07-21 Accepted:2026-02-02 Online:2026-03-01 Published:2026-03-06
  • Contact: LI ShaoKun

Abstract:

【Objective】 This study systematically quantified the contribution rates and spatial heterogeneity of protein, starch, and fat—the three major nutritional components—to maize kernel test weight formation, and elucidated how genetic background, ecological region, and cultivation density interactively modulate both nutritional quality and test weight. The findings aim to establish a science-based foundation for region-specific optimization of maize quality and to advance an integrated “high-yield-high-quality- high-efficiency” production paradigm.【Method】 A nationwide field survey was conducted across four major maize-producing regions in China, encompassing 718 representative kernel samples from 77 leading cultivars grown under 24 distinct planting density gradients (37 500-127 500 plants/hm2). All samples were naturally air-dried to standardized moisture content (14% w.b.) prior to uniform physicochemical analysis. Protein, starch, and fat contents were determined using calibrated near-infrared reflectance spectroscopy (NIRS), and test weight was measured with a certified grain test weight instrument (ISO 7971-3 compliant). To dissect the complex determinants of test weight, we implemented a hierarchical analytical framework integrating: (i) multiple linear regression to estimate independent linear effects; (ii) random forest modeling to capture nonlinear interactions and relative feature importance; and (iii) structural equation modeling (SEM) to infer directional causal pathways among traits. Three-way ANOVA was further employed to assess the main and interactive effects of cultivar, ecological region, and cultivation density on test weight and each nutritional component.【Result】 Protein (β=8.406, P<0.001) and starch (β=6.413, P<0.001) emerged as statistically robust and biologically dominant drivers of test weight, accounting for 28% and 45% of the total explained variance in the random forest model, respectively—both exhibiting high path coefficient stability in SEM (standardized coefficients ≥0.72, P<0.001). In contrast, fat showed negligible explanatory power (2%), and its effect failed to reach statistical significance (P=0.09). Three-way ANOVA confirmed highly significant (P<0.001) main effects and two- and three-way interactions among cultivar, ecological region, and density for test weight, protein, and starch—indicating strong contextual dependency. Spatially, protein contributed most strongly in the Northeast spring maize region (43.9% of model variance), whereas starch dominated in the Huang-Huai-Hai summer maize region (52.9%). Critically, the synergistic contribution of protein and starch jointly explained 81.0% and 85.0% of the total model variance in these two regions, respectively. Structural equation modeling revealed a direct positive effect of protein on test weight, but an indirect negative effect stemming from the compensatory relationship between protein and starch accumulation, which underscores the physiological trade-off in kernel sink-filling.【Conclusion】 Maize test weight formation was a biologically synergistic process driven by protein and starch, with fat playing no substantial role. Significant interactions existed among cultivar, ecological region, and density, with the same cultivar exhibiting distinct regulatory pathways under different ecological and cultivation conditions. Consequently, the Northeast region should prioritize high-protein cultivar selection and precise nitrogen management, while the Huang-Huai-Hai region should enhance carbon assimilation efficiency and regulate key starch-synthesis enzymes. All production areas should achieve a precise "cultivar-region-practice" matching strategy to synergistically improve maize yield and quality.

Key words: maize, nutritional quality, kernel test weight, regional heterogeneity, multi-model analysis

Table 1

Overview of national maize sampling sites"

玉米产区
Maize production region
省份
Province
样品数量
Number of samples
取样地区
Sampling region
播种密度
Sowing density (plants/hm2)
黄淮海夏玉米区
Huang-Huai-Hai summer maize region
安徽
Anhui
34 阜阳Fuyang(17)、宿州Suzhou(7)、淮北Huaibei(3)、亳州Bozhou(4)、蚌埠Bengbu(3) 82500、85500、90000、97500、105000
河北
Hebei
34 石家庄Shijiazhuang(25)、邯郸Handan(4)、邢台Xingtai(5) 60000、67500、75000、82500
河南
Henan
47 焦作Jiaozuo(4)、商丘Shangqiu(4)、许昌Xuchang(4)、驻马店Zhumadian(3)、开封Kaifeng(4)、新乡Xinxiang(4)、三门峡Sanmenxia(3)、安阳Anyang(3)、鹤壁Hebi(2)、平顶山Pingdingshan(5)、周口Zhoukou(8)、洛阳Luoyang(3) 66000、67500、75000、82500、90000、105000
江苏
Jiangsu
17 盐城Yancheng(5)、徐州Xuzhou(8)、连云港Lianyungang(4) 55500、64500、67500、75000、82500、90000
山东
Shandong
108 青岛Qingdao(10)、烟台Yantai(10)、潍坊Weifang(10)、济宁Jining(22)、枣庄Zaozhuang(5)、菏泽Heze(5)、聊城Liaocheng(21)、德州Dezhou(20)、淄博Zibo(5) 63000、67500、72000、75000、82500、85500、87000、90000、93000、97500
陕西
Shaanxi
50 榆林Yulin(25)、西安Xi'an(6)、渭南Weinan(6)、宝鸡Baoji(7)、咸阳Xianyang(6) 63000、67500、72000、75000、82500、87000、90000
西北春玉米区
Northwest spring maize region
甘肃Gansu 15 平凉Pingliang(10)、白银Baiyin(5) 60000、64500、67500、75000
内蒙古
Inner Mongolia
16 呼和浩特Hohhot(5)、巴彦淖尔Bayannur(5)、包头Baotou(3)、鄂尔多斯Ordos(3) 67500、75000、82500、85500
新疆Xinjiang 15 昌吉Changji(5)、博乐Bole(5)、乌鲁木齐Urumqi(5) 112500、120000、127500
山西Shanxi 22 长治Changzhi(18)、晋城Jincheng(4) 67500、75000
西南夏玉米区
Southwest summer maize region
广西
Guangxi
20 崇左Chongzuo(5)、南宁Nanning(5)、贵港Guigang(4)、河池Hechi(4)、百色Baise(2) 42000、45000、52500
湖北Hubei 16 恩施Enshi(12)、宜昌Yichang(4) 37500、40500、45000、75000
四川Sichuan 10 绵阳Mianyang(5)、德阳Deyang(5) 52500、57000、63000、90000
东北春玉米区
Northeast spring maize region
黑龙江
Heilongjiang
162 黑河Heihe(37)、齐齐哈尔Qiqihar(44)、鹤岗Hegang(2)、双鸭山Shuangyashan(6)、绥化Suihua(19)、伊春Yichun(6)、哈尔滨Harbin(8)、佳木斯Jiamusi(8)、大庆Daqing(32) 60000、67500、75000、82500、90000、97500、105000
吉林
Jilin
69 长春Changchun(4)、四平Siping(2)、白城Baicheng(8)、松原Songyuan(12)、吉林Jilin(40)、敦化Dunhua(3) 52500、55500、60000、67500、72000、75000、82500、90000
内蒙古
Inner Mongolia
29 通辽Tongliao(14)、赤峰Chifeng(15) 60000、72000、75000、82500、90000、97500、112500
辽宁
Liaoning
60 大连Dalian(5)、丹东Dandong(7)、营口Yingkou(5)、阜新Fuxin(22)、沈阳Shenyang(10)、铁岭Tieling(11) 60000、67500、75000、82500、97500

Table 2

Key indicators of the four major maize production regions"

产区
Region
蛋白质
Protein
(%)
脂肪
Fat
(%)
淀粉
Starch
(%)
容重
Test weight
(g·L-1)
平均种植密度
Average planting density (plants/hm2)
东北春玉米区 Northeast spring maize 10.08±0.83a 3.80±0.43c 74.37±1.13b 775.06±25.98a 77944
西北春玉米区 Northwest spring maize 9.17±0.84b 3.98±0.34b 74.55±0.85a 771.23±22.69a 83077
西南夏玉米区 Southwest summer maize 9.66±0.84ab 4.80±0.39a 73.68±0.78c 756.27±23.64c 54545
黄淮海夏玉米区 Huang-Huai-Hai summer maize 9.35±0.81b 3.97±0.69b 75.43±3.82a 769.42±22.36b 79382

Table 3

Multiple linear regression coefficients for national maize"

变量
Variables
非标准化回归系数
β (Coefficient)
标准化回归系数
β (Standardized)
标准误
SE
t值
t value
P
P value
95%置信区间
95% CI
蛋白质Protein 6.41 0.33 1.61 3.99 P<0.001 3.25-9.57
脂肪Fat 5.93 0.12 3.51 1.69 P =0.09 -0.95-12.81
淀粉Starch 8.41 0.31 2.22 3.78 P<0.001 4.06-12.76

Fig. 1

Path diagram of the structural equation model (SEM) A: Path diagram of structural equation model for nationwide maize; B: Path diagram of structural equation model for spring maize in northeast region; C: Path diagram of structural equation model for summer maize in Huang-Huai-Hai region"

Table 4

Statistical test results of structural equation modeling path coefficients for nationwide maize"

路径关系
Pathway relationship
非标准化系数
Unstandardized coefficient
标准化系数
Standardized coefficient
标准误
SE
t值
t value
P
P value
蛋白质→容重 Protein→test weight 8.41 0.62 1.40 6.02 P<0.001
淀粉→容重 Starch→test weigh 6.41 0.54 1.09 5.87 P<0.001
蛋白质→淀粉 Protein→starch -0.10 -0.18 0.05 -2.11 P=0.035

Table 5

Parameter optimization for the national maize test weight model"

参数组合
Parameter combination
决定系数
R2
均方根误差
RMSE (g·L-1)
平均绝对误差
MAE (g·L-1)
调整后决定系数Adjusted R2 性能变化
Performance change
ntree=500, mtry=1, max_depth=6 0.589 7.128 5.812 0.587 欠拟合Underfitting
ntree=500, mtry=1, max_depth=8 0.604 6.981 5.642 0.602 优化中Optimizing
ntree=500, mtry=1, max_depth=10 0.597 7.051 5.735 0.595 基准模型Benchmark
ntree=800, mtry=1, max_depth=6 0.633 6.712 5.423 0.631 -
ntree=800, mtry=1, max_depth=8 0.648 6.573 5.281 0.646 +8.5%▲
ntree=800, mtry=1, max_depth=10 0.642 6.650 5.432 0.640 区域最佳候选Regional best candidate
ntree=1100, mtry=1, max_depth=6 0.638 6.682 5.301 0.636 -
ntree=1100, mtry=1, max_depth=8 0.659 6.501 5.152 0.657 +10.4%▲
ntree=1100, mtry=1, max_depth=10 0.645 6.619 5.231 0.643 全国最优模型National optimal

Table 6

Comparison of standardized values for nutritional component contributions across three models"

模型类型 Model type 淀粉贡献 Starch contribution 蛋白质贡献 Protein contribution 脂肪贡献 Fat contribution
多元线性回归 Multiple linear regression 0.31 0.33 0.12
随机森林模型 Random forest model 0.45 0.28 0.02
结构方程模型 Structural equation model 0.64 0.84 0.03

Fig. 2

Comparison of variables in random forest models of corn nutritional quality for two major maize production regions"

Table 7

Multiple linear regression coefficients for two major maize production regions"

玉米产区
Maize production region
变量
Variables
非标准化回归系数
β (Coefficient)
标准化回归系数
β (Standardized)
标准误
SE
t值
t value
P
P value
95%置信区间
95% CI
黄淮海夏玉米区
Huang-Huai-Hai summer maize region
蛋白质Protein 12.67 0.19 6.41 1.98 0.05 0.11-25.23
脂肪Fat 1.60 0.02 6.52 0.25 0.81 -11.18-14.38
淀粉Starch 8.92 0.26 3.36 2.66 0.01 2.33-15.51
东北春玉米区
Northeast spring maize region
蛋白质Protein 18.85 0.44 3.21 5.87 0.00 12.56-25.14
脂肪Fat 6.31 0.08 4.95 1.27 0.20 -3.38-16.00
淀粉Starch 11.36 0.37 2.35 4.82 0.00 6.75-15.97

Table 8

Three-way ANOVA results for the effects of variety, region and planting density on kernel test weight and nutritional qualities"

分析因素
Analysis of factors
容重 Test weight 蛋白质 Protein 淀粉 Starch
F值 F value PP value F值 F value
品种 Variety 38.2 P<0.001 35.6
区域 Region 42.5 P<0.001 28.9
密度 Density 6.9 P<0.05 3.2
品种×区域 Variety×region 15.3 P<0.001 12.8
品种×密度 Variety×density 3.1 P<0.05 2.8
区域×密度 Region×density 7.2 P<0.01 5.6
品种×区域×密度 Variety×region×density 4.2 P<0.05 3.3
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