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Article

Effects of Nitrogen Fertilizer Application on the Lodging Resistance Traits, Yield, and Quality of Two Gluten Types of Wheat

by
Xueling Hu
1,
Peiyu Tian
2,
Wen Fu
3,
Zhihao Tian
4,
Mengdi Du
1,
Zhishang Chang
1,
Youliang Ye
1,
Xiangping Meng
1,* and
Yang Wang
1
1
College of Resources and Environment, Henan Agricultural University, Zhengzhou 450002, China
2
College of Resources and Environment, China Agricultural University, Beijing 100193, China
3
Xuchang Agricultural Technology Extension Station, Xuchang 452570, China
4
Yuzhou Agricultural Technology Extension Station, Xuchang 452570, China
*
Author to whom correspondence should be addressed.
Agriculture 2025, 15(6), 637; https://doi.org/10.3390/agriculture15060637
Submission received: 11 February 2025 / Revised: 11 March 2025 / Accepted: 14 March 2025 / Published: 18 March 2025
(This article belongs to the Section Crop Production)
Browse Figures
Versions Notes

Abstract

:
The Huang–Huai–Hai Plain is a primary wheat production base in China, where lodging remains a critical constraint limiting yield improvement and quality enhancement. Both nitrogen application and wheat varieties are key factors influencing crop lodging resistance. This study aimed to comparatively analyze the differential responses of wheat varieties with distinct gluten properties to nitrogen fertilization gradients and elucidated the physiological mechanisms underlying the nitrogen-mediated regulation of lodging resistance in gluten-type wheat. A two-year field experiment was conducted in Xuchang City, Henan Province, from 2019 to 2021. The experimental design incorporated four varieties of wheat (two medium-gluten wheat varieties, YM49-198 and JM325, and two strong-gluten wheat varieties, XN979 and JM44) and five nitrogen (N) fertilizer levels: 0 kg·ha−1 (N0), 120 kg·ha−1 (N120), 180 kg·ha−1 (N180), 240 kg·ha−1 (N240), and 360 kg·ha−1 (N360). Each treatment was repeated three times, and each plot was completely randomly arranged in the field. An appropriate amount of nitrogen fertilizer significantly increased the wheat yield, with the 240 kg ha−1 treatment achieving maximum yields for YM49-198, JM325, and JM44 from 2020 to 2021, but not for XN979. Quality parameters were significantly affected by varieties and nitrogen fertilizer levels. The results showed that the crude protein contents of XN979 and JM44 were 15.13% and 18.06%, respectively, under the N240 treatment; the lodging resistance index of the medium-gluten wheat was higher than that of the strong-gluten wheat. Under the N240 treatment in 2020–2021, the lodging resistance indexes of YM49-198, JM325, XN979, and JM44 were 12.2, 13.9, 7.9, and 11.7, respectively. Nitrogen fertilizer can increase wheat yield and ensure quality, but excessive application can decrease these factors and intensify lodging risk. The lodging index of the medium-gluten wheat was more sensitive to the amount of nitrogen fertilizer. When the nitrogen application is 240 kg·ha−1, the quality indicators of medium- and strong-gluten wheat should meet standards, and the yield will be stable in the Huang–Huai–Hai Plain. These findings highlight the importance of adopting precision nitrogen management strategies and gluten-type-specific cultivation practices in wheat production systems. This could effectively balance yield stability, quality optimization, and lodging risk mitigation to ensure the sustainable intensification of wheat cultivation in the Huang–Huai–Hai Plain and similar agro-ecological regions.

1. Introduction

As a staple cereal crop, wheat plays a crucial role in global food security. As the world’s largest wheat producer, China contributes significantly to this agricultural sector [1]. The Huang–Huai–Hai Plain is the core production hub for wheat in China, contributing approximately 60% of the nation’s total wheat output. Its production fluctuations directly affect the national Consumer Price Index (CPI) for grain by 0.8–1.2 percentage points (China Food Security Wheat Paper) [2]. Nevertheless, lodging remains the primary constraint limiting yield enhancement and quality improvement in wheat cultivation in this area [3]. Lodging adversely impacts photosynthetic efficiency, disrupts nutrient–water assimilation, and ultimately reduces grain output by 20–50% [4]. Therefore, enhancing lodging resistance is critical for optimizing both productivity and end-use quality, particularly to meet escalating demands for premium wheat products amid rising living standards in China [5].
Genetic diversification strategies have proven effective in reconciling lodging resistance with yield potential [6]. Plant breeders continually introduce genes to produce high-quality wheat varieties with strong lodging resistance [7]. Khan, Hanif et al. [8] reduced the incidence of lodging in high-yielding varieties by spraying chlormequat chloride growth regulators. Plant height has a strong and direct impact on lodging resistance because it affects the bending stress experienced in the plant stem [9]. However, shorter varieties can be grown with more inputs, such as fertilization and irrigation, because they may not lodge as much as taller varieties [10]. Lodging can also cause other problems, such as decreased grain quality, reduced harvesting efficiency, and increased labor costs. The lodging resistance of different wheat genotypes varies, and most quality traits are much more affected by environmental factors than by genotypes [11]. With the development of the economy, the demand for special flours for high-quality flour-based foods in China is increasing [12]. To improve wheat, it is necessary to select high-quality varieties and improve their lodging resistance.
Nitrogen management constitutes a pivotal factor in wheat production systems, governing both metabolic processes and structural integrity [13,14]. While judicious nitrogen application enhances yield through tiller promotion and photosynthetic optimization [15,16,17,18], excessive inputs exacerbate lodging risks by elevating plant height, basal internode elongation, and the center of gravity [19]. Conversely, nitrogen reduction strategies improve stem robustness through increased culm diameter and wall thickness, albeit potentially compromising grain protein content [20]. This creates a critical trade-off between yield quantity and quality parameters. Research has shown a significant correlation between lodging resistance and anatomical features, including mechanical tissue, the number of vascular bundles, the area of a single vascular bundle, and pith diameter [21]. Meanwhile, the application rate of nitrogen fertilizer can affect the lodging resistance of crops by influencing the structural composition and cross-sectional architecture of plant stems. From a quality perspective, nitrogen is an important factor in the protein and wet gluten content in wheat, and the quality of wheat dough and pasta is largely influenced by the protein content and gluten quality [22,23,24,25]. Nitrogen fertilizer plays a key role in increasing the protein content and overall grain quality [26,27], and grain protein content increases with increasing nitrogen applications [28]. Compared with medium-gluten wheat, increasing nitrogen fertilizer can significantly increase the activity of key enzymes for protein synthesis in strong-gluten wheat, thereby improving its quality [29]. However, this also makes high-gluten wheat more prone to lodging. Under the same nitrogen form, the protein content of each component in the grains of strong-gluten wheat is higher than that of medium-gluten wheat, but the grain yield is lower [30].
In this study, we compared differences in the response of different gluten wheat types to N fertilizer dosages. It could also provide a scientific basis for developing a reasonable fertilizer application strategy to ensure stable wheat yield and the safety of grain silos in the Huang–Huai–Hai Plain. We hypothesized that (1) nitrogen application significantly influences the lodging resistance due to their distinct genotypic characteristics; (2) nitrogen application plays a critical role in determining the crude protein and total protein content of wheat grains; (3) a trade-off exists between yield, quality, and lodging risk in different wheat types under varying nitrogen regimes.

2. Materials and Methods

2.1. Experimental Site

A field experiment was conducted from mid-October 2019 to mid-June 2021 at the experimental farm of Henan Agricultural University, Xuchang, China (34°16′ N, 113°21′ E). The area has a warm temperate climate characterized by semi-humidity and continental monsoonal influences. Daily precipitation and average daily temperatures during winter wheat fertility in 2020 and 2021 are shown in Figure 1. The wheat-growing seasons of 2019–2020 and 2020–2021 exhibited marked differences in total rainfall and its temporal distribution. The 2019–2020 season received 139.9 mm of rainfall, lower than the 201.3 mm recorded in the 2020–2021 season, representing a 30.5% reduction. Furthermore, the distribution of rainfall during key growth stages showed contrasting patterns between the two years. In 2019–2020, the leaf emergence and tillering stages received only 2.1 mm and 66.0 mm of rainfall, respectively (Figure 1A). Conversely, during the 2020–2021, these stages experienced substantially higher and lower rainfall amounts at 55.4 mm and 9.5 mm, respectively (Figure 1B). The upper 20 cm at the experimental site was tidal brown soil with the following properties: pH 8.2, 16.3 g kg−1 total organic matter, 1.04 g kg−1 total nitrogen, 20.0 mg kg−1 available phosphorus, and 113.7 mg kg−1 available potassium [31]. Soil property data were collected each year and averaged across the two years.

2.2. Experimental Design

This experiment had a two-factor design with two wheat gluten types (medium-gluten and strong-gluten) and different nitrogen applications. The medium-gluten varieties were Yumai49-198 (YM49-198) and Jimai324 (JM325), and the strong-gluten cultivars were Xinong979 (XN979) and Jimai44 (JM44). The five nitrogen applications were 0 kg·ha−1 (N0), 120 kg·ha−1 (N120), 180 kg·ha−1 (N180), 240 kg·ha−1 (N240), and 360 kg·ha−1 (N360). Twenty treatments were applied, and each included three replicates, with each plot measuring 30 m² (5 m × 6 m).
Regarding the four wheat varieties, YM49-198 had a capacity of 790~810 g/L, 14.74% crude protein content, 32.5% wet gluten content, and a 38 mL sedimentation value; JM325 had a capacity of 802 g/L, 14.36% crude protein content, 30.1% wet gluten content, and a 24.5 mL settling value; XN979 had 14–16% crude protein content, 32%~34% wet gluten content, and a 53.2 mL sedimentation value; JM44 had 15.35% crude protein content, 35.1% wet gluten, and a 51.5 mL sedimentation value [32,33,34,35].
Before wheat sowing, 50% of the prescribed nitrogen dose, with 90 kg of P2O5 ha−1 and 90 kg of K2O ha−1, was applied as basal fertilizer. The other half of the nitrogen was applied at the jointing stage as a top dressing. The previous crop at the trial site was summer maize, and the straw was removed. Wheat was sown mechanically at a seeding rate of 750 kg·ha−1 in both years, with sowing dates on 20 October 2019, and 23 October 2020, and corresponding harvest dates on 1 June 2020, and 5 June 2021. Chemical controls were judiciously applied throughout the cropping period to effectively manage weeds, diseases, and pests.

2.3. Measurement Index and Methods

2.3.1. Measurement of Grain Yield and Yield Components

The plants were sampled diagonally from two rows per meter to determine yield components during the ripening period. After counting the total spike number to determine the number of spikes per square meter, the plant samples were separated into stems plus leaves and spikes. All the spikes were hand-threshed and cleaned, and the total grain number and grain weight were recorded after oven-drying at 70 °C when they reached a constant weight. Grain number per spike and spike numbers were calculated. Grain yield was determined by mechanically harvesting the central 0.667 m2 area.

2.3.2. Number of Productive Tillers

Before tillering at the wheat seedling stage, we selected two adjacent representative rows with a length of 1 m in each plot for fixed-point observation, counting the number of basic seedlings in the two rows. The individual stages of growth were determined according to the BBCH scale (Biologische Bundesanstalt, Bundessortenamt und CHemische Industry): leaf development (L)—BBCH 10–13; tillering (T)—BBCH 20–29; stem elongation (S)—BBCH 30–39; flowering (F)—BBCH 61–69; and ripening (R)—BBCH 83–89 (Figures S1 and S2).

2.3.3. Measurement of Vascular Bundle Number, Area, and Mechanical Tissue Thickness

At the grain filling stage, ten plants per plot were earmarked. Their second internodes from the basal stems were collected and fixed in a fixative solution of formyl aceto-alcohol (FAA) (50% ethanol, 10% formalin, 5% glacial acetic acid, and 35% double distilled water). A series of preservation techniques yielded permanent paraffin sections: dehydration, clearing, wax infiltration, embedding, slicing, mounting, dewaxing, staining with saffron and fast green, and sealing. The sections were photographed using an electron microscope (Leica DM750, Leica Microsystems Wetzlar GmbH, Hessen, Germany) to observe the number of large vascular bundles (inner-ring vascular bundles) and small vascular bundles (outer-ring vascular bundles) in parenchyma tissues, as well as the thickness of the mechanical tissue and cell walls. The ImageJ software (1.54) was used to measure the areas of large and small vascular bundles, the thickness of the mechanical tissue, and cell wall width. Each material was observed 10 times, and the average values were taken [36].

2.3.4. Investigation of Lodging-Related Traits

Lodging-related traits were determined at the fruit development stage (BBCH 70–82). Ten representative single stems with consistent growth were selected from each plot. The morphological parameters of the whole plant measured included height, the center of gravity height (from the plant base to the fulcrum of the balance location), and the length of each internode. Stem diameter was obtained by measuring the diameter of the second internode. Because lodging usually occurs in basal internodes, the second internode at the base of the stem was taken when the leaf sheath was stripped, and the stem breaking strength against collapse was determined using a YYD-1 stalk strength instrument (Zhejiang Top Instrument Co., Ltd., Hangzhou, China). The lodging resistance index was calculated according to the following formula: lodging resistance index = breaking resistance strength/center of gravity height [37].

2.3.5. Measurement of Grain Quality

The wheat seeds were ground in a lap mill (Chopin CD1, Henan Rongcheng United Technologies Co., Ltd., Zhengzhou, China). The quality indicators included the crude protein content (%), wet gluten values (%), Zeleny sedimentation (mL), flour extraction rate (%), and bulk density (g/L). These parameters were determined following GB/T17320-2013 [38], and the starch (%) was determined using the Ewers polarimetric method [39]. All determinations were performed in triplicate.

2.3.6. Measurement of Stem Nutrient Elements

The twenty representative plants were chosen at the fruit development stage. The second internode was taken from each plant, with the leaves and leaf sheaths stripped off. The samples were dried at 105 °C for 30 min and then at 80 °C to a constant weight, and they were then ground to pass through a 1 mm mesh screen. After digestion using a microwave digestion instrument (MARS6, PYNN Corp., Matthews, NC, USA), the stem nutrient element contents (K, Ca, Mg, and Fe) were determined using an ICP-OES instrument (Optima 8000, Shanghai Shanfu Electronic Technology Co., Ltd., Shanghai, China) [40].

2.4. Statistical Analysis

The analyses of variance of yield-related parameters, lodging-related parameters, wheat stem structure-related parameters, and stem nutrient elements were performed with general linear modeling (GLM) in SPSS 25.0 (SPSS, Inc., Chicago, IL, USA). All data were analyzed with a two-way analysis of variance (ANOVA) using the least significant difference (LSD) multiple comparisons test (p < 0.05). Figures were generated using Origin Pro (2021).

3. Results

3.1. Vascular Bundles of Stalk

With increasing nitrogen application rates, both large and small vascular bundle numbers exhibited significant decreasing trends across all four wheat cultivars (p < 0.05), though cultivar-specific variations were observed (Figure 2A,C). Notably, strong-gluten cultivars demonstrated 2.4% and 21.6% higher in large and small vascular bundles, respectively, compared to medium-gluten cultivars. XN979 displayed the highest small vascular bundle amount (22.47) among the tested cultivars. The opposite response to the nitrogen fertilizer emerged in the big and small vascular bundles (Figure 2B,D). While increasing nitrogen levels reduced small vascular bundle area and increased the large vascular bundle area in the four wheat variations, the differences in the mechanical tissue thickness of different varieties were relatively small, and the change trends in the varieties were not consistent at different nitrogen application levels. YM49-198, JM325, and XN979 reached maximum thickness values at N180 (91.79, 97.70, and 102.76 mm, respectively), whereas JM44 achieved peak thickness (104.82 mm) at N240.

3.2. Mineral Nutrient Element Contents in Wheat Stalks

Nitrogen application significantly affected the elemental K, Ca, Mg, and Fe contents in wheat stalks (Figure 3). With the increase in nitrogen application, the contents of these four elements showed a clear upward trend, but the variation patterns among different varieties were different. The K content from XN979 and JM44 was highest and lowest, respectively. Compared with the N0 treatment, the K content in the stalk of the two gluten wheat varieties under the N360 treatment increased by 45.0% (YM49-198), 58.0% (JM325), 28.1% (XN979), and 26.0% (JM44), respectively. For elemental Ca, the Ca content of different strong-gluten wheat and medium-gluten wheat varieties differed, as shown by the fact that YM49-198 > JM325, XN979 > JM44; Mg content in stalks of strong-gluten varieties increased significantly when N application was more than 180 kg ha−1, with an average increase of 35.35% (XN979) and 70.75% (JM44) compared with treatments lower than 180 kg·ha−1. Overall, the interaction of N application and variety significantly affected K, Ca, Mg, and Fe contents’ concentrations in the stalks.

3.3. Lodging Resistance Index and Its Related Parameters

The nitrogen application significantly increased the plant height and center of gravity height of both types of wheat, with the heights recorded in 2020–2021 higher than those in 2019–2020 for all varieties (Table 1 and Table 2). With increasing nitrogen applications, both plant height and center of gravity height exhibited a trend of first increasing, decreasing, and increasing again in 2020–2021. Compared with the N0 treatment, the medium-gluten wheat (JM325) under N240 treatment showed increases of 37.4% in plant height and 33.6% in center of gravity height. The interaction of variety and nitrogen significantly affected the plant height and center of gravity height of the two types of wheat.
Table 1 and Table 2 showed that the interaction between variety and N application significantly affected the stem breaking strength and lodging resistance index of the wheat. The stem breaking strength was higher in both wheat types at N0 and tended to increase and then decrease with increasing N applications, reaching a minimum at N360. The decline in stem breaking strength was greater in the medium-gluten wheat than in the strong-gluten wheat. Over two years, the average stem breaking strength of the medium-gluten wheat exceeded that of the strong-gluten wheat by 31.7% (2019–2020) and 25.8% (2020–2021), indicating that nitrogen applications had a more pronounced effect on the flexural strength of the medium-grain wheat. The lodging resistance index of both wheat types decreased significantly with increased nitrogen. In 2020–2021, the lodging resistance index of the medium-gluten wheat was 27.0% higher than that of the strong-gluten wheat, indicating that the medium-gluten wheat had better bending resistance.

3.4. Grain Yield

In two years, the yield of medium-gluten wheat was higher than strong-gluten wheat (Figure 4A,B). But the strong-gluten wheat yield in 2020–2021 was significantly lower than in 2019–2020. With the increased nitrogen application, the yield of the two types of gluten wheat showed a trend of initially increasing and then slightly decreasing, remaining at a high level between N180 and N360. Compared with N0 treatment, JM325 (under N240 treatment) and XN979 (under N360 treatment) increased the yield by 169.7% and 67.5%, respectively, in 2020–2021.
With the increased nitrogen application, the thousand-grain weight of both wheat types decreased, remaining at a high level between the N0 and N180 treatments. The overall thousand-grain weight of the medium-gluten wheat was higher than that of the strong-gluten wheat in 2020–2021 (Figure S3C,F).

3.5. Grain Qualities

The nitrogen application significantly affected the crude protein, wet gluten, starch, sedimentation value, flour yield, and bulk density of the different gluten wheat types (Figure 5). In the medium-gluten wheat varieties, the nitrogen application significantly enhanced grain quality parameters. Under N240 treatment, crude protein content and wet gluten content increased by an average of 35.17% and 44.64% in YM19-198 and 26.42% and 27.33% in JM325, respectively, compared with the N0 treatment. These improvements were amplified under the N360 treatment, with YM19-198 reaching 38.77% and 50.57% and JM325 achieving 31.77% and 33.14% for crude protein and wet gluten content, respectively. The strong-gluten wheat varieties exhibited distinct qualities compared with the medium-gluten wheat. XN979 and JM44 showed significant differences in starch content, flour extraction, and bulk density. Notably, the sedimentation values of JM44 were 52.32% and 55.95% higher than those of YM19-198 and JM325, respectively, under the N240 treatment, highlighting the superior gluten strength of the strong-gluten wheat.

3.6. Analysis of Structural Equation Modeling

In this research, smart PLS 3 structural equation modeling was applied, showing that nitrogen fertilization significantly affected yield, quality, and stem bending resistance in both medium-gluten wheat (YM49-198 and JM325) and strong-gluten wheat (XN979 and JM44) (Figure 6 and Figure 7). For medium-gluten wheat, applied nitrogen was significantly and positively correlated with stem thickness, yield, and quality and negatively correlated with vascular structure; the path coefficients were 0.400, 0.507, 0.930, and −0.838, respectively. The vascular structure significantly affected stem bending resistance and, thus, yield formation, and applying N fertilizer significantly affected stem bending resistance and yield. In the strong-gluten wheat, nitrogen application was negatively correlated with stem thickness and vascular bundle structure and significantly positively correlated with yield and quality. Finally, stem bending resistance significantly affected the yield and quality of wheat.

4. Discussion

4.1. Effect of Nitrogen and Cultivar on Lodging Resistance in Winter Wheat

In our study, nitrogen (N) application and cultivar significantly influenced lodging resistance, primarily through modifications to stem anatomical traits and mechanical properties. The medium-gluten wheat varieties (e.g., JM325) exhibited superior bending resistance to the strong-gluten wheat varieties (e.g., XN979), with the average stem breaking strength values of 46.6–67.9% being higher across all N treatments (Table 1 and Table 2). This divergence likely stems from cultivar-specific vascular bundle organization: the medium-gluten wheat maintained denser small vascular bundles (Figure 2A,C), which enhance mechanical load distribution and stem flexural rigidity [41] Structural equation modeling (SEM) revealed that nitrogen application negatively impacted vascular bundle structure (path coefficient: −0.838) in the medium-gluten wheat, yet its robust stem anatomy compensated by maintaining higher bending resistance (Figure 6 and Figure 7). Excessive nitrogen fertilizer application increases plant height and raises the center of gravity across all straw internodes. Concurrently, it reduces stem diameter. These structural changes collectively weaken the plant’s lodging resistance [22,24]. Our experimental results exhibited a consistent pattern in this regard. Notably, the strong-gluten wheat displayed greater susceptibility to nitrogen-induced lodging, particularly under the N360 treatment regime, where stem breaking strength declined by 76.8% in XN979 (Table 1). This may be due to the difference in genotype, which makes strong-gluten wheat more sensitive to nitrogen fertilizer than medium-gluten wheat; excessive N in strong-gluten wheat is more likely to promote cell elongation over wall thickening, compromising structural integrity [42]. Furthermore, many studies have demonstrated that nitrogen fertilization timing and methods substantially influence cell wall development. Specifically, these practices regulate both the formation of cellulose microfibrils and the spatial organization dynamics of microtubules, which collectively govern cellulose biosynthesis and ultimately affect plant lodging resistance [43,44,45].

4.2. Effects of Nitrogen and Cultivar on Grain Yield

The yield of wheat was determined by three factors [46]. With the increase in nitrogen applications, the number of tillers (Figures S1 and S2), the number of grains per spike (Figure S3B,E), and the yield of the two types of gluten wheat all showed a gradually increasing trend in our research. Overall, the medium-gluten wheat yield was higher than that of the strong-gluten wheat; the average growth rates of the medium-gluten and strong-gluten wheat were 74% and 94% in 2019–2020 and 62% and 75% in 2020–2021, respectively. This indicated that wheat yield responses to N fertilization exhibited gluten-type-specific patterns, with the medium-gluten wheat achieving higher stability across N regimes. JM325 produced maximum yields under N240, which was more than N0 by 100.2%, whereas the strong-gluten wheat (XN979) peaked earlier at N360 (Figure 4B). This divergence may reflect the contrasting N utilization strategies of the varieties. Strong-gluten wheat may be more efficient in nitrogen utilization because of its complex gluten matrix structure. Under equivalent nitrogen inputs, these cultivars preferentially allocate nitrogen resources toward gluten and storage protein biosynthesis. Conversely, medium-gluten wheat exhibits reduced nitrogen assimilation capacity, consequently requiring lower nitrogen inputs to achieve optimal growth performance [47]. Our experimental findings corroborate this view; under the N240 treatment, the medium-gluten wheat demonstrated significantly greater stem diameter and a higher ear number (Figure S3A,D) than the strong-gluten wheat.

4.3. Effects of Nitrogen and Cultivar on Grain Quality

The results of this experiment showed that the crude protein, wet gluten, starch, and sedimentation value of the two types of gluten wheat increased with increasing nitrogen applications. Fu [30] found a similar result in Shandong Province. However, our result also showed that the same nitrogen application differentially modulated grain quality parameters between gluten types, with strong-gluten wheat (XN979 and JM44) achieving superior protein and gluten content under elevated N regimes. Under N240, crude protein and wet gluten in XN979 increased by 40.1% and 41.5%, respectively, meeting GB/T17320-2013 [48] national standards (Figure 5A,B). This enhancement likely stems from nitrogen-rich fertilization supplying abundant nitrogen substrates, which upregulates key protein synthesis pathway enzymes, including glutamine synthetase (GS) and glutamate–pyruvate aminotransferase (GOGAT) [29]. Additionally, elevated nitrogen inputs significantly increase storage proteins (globulins and wheat albumins) in strong-gluten wheat, potentially contributing to its superior quality indices [49]. Notably, water conditions, such as irrigation and rainfall, also modulate wheat quality parameters, as demonstrated by parallel studies [49,50].

5. Conclusions

We systematically analyzed the effects of nitrogen application on the lodging, yield and quality of different wheat varieties in the Huang–Huai–Hai Plain through a two-year field experiment. Results demonstrated that nitrogen application could improve wheat yield, increase wheat’s ability to resist lodging and improve grain quality, but excessive application would increase plant height and center of gravity height, and exacerbate the risk of falling. Compared with strong-gluten wheat, medium-gluten wheat was more susceptible to fall when N application exceeded 240 kg·ha−1. Regardless of strong or medium-gluten wheat, the yields under nitrogen application 180–240 kg·ha−1 showed a trend of rapid increase followed by slow increase with increasing N application, and the average yield of medium-gluten wheat under the same fertility treatment was significantly higher than that of strong-gluten wheat in both years. The highest grain quality indexes were observed in the N240 treatments for strong-gluten wheat (XN979, JM44), but excessive N application (>240 kg·ha−1) resulted in a multiplication of the risk of lodging. Therefore, in order to balance the risk of lodging, yield and grain quality in the Huang–Huai–Hai Plain, the recommended N application rate is 180–240 kg·ha−1 for medium-grain wheat and not more than 240 kg·ha−1 for strong-grain wheat.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agriculture15060637/s1, Figure S1: Effects of nitrogen application rate on population of two gluten types of wheat (2019–2020); Figure S2: Effects of nitrogen application rate on population of two gluten types of wheat (2020–2021); Figure S3: Effects of nitrogen application rate on yield components of two types of gluten wheat. A, B and C represent the spike number per m2, grain number per spike and the thousand-grain weight in 2019–2020, respectively; D, E and F represent the spike number per m2, grain number per spike and the thousand-grain weight in 2020–2021, respectively.

Author Contributions

X.H. and P.T. conducted literature search and data collection, and prepared visual presentations; W.F. and Z.T. participated in data collection; M.D. and Z.C. participated in literature search and prepared visual presentations; Y.W. and X.M. conceptualized the work, performed administration of the projects and reviewed different versions of the paper; Y.Y. provided ideas and participated in revising the article. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Key Research and Development Program of China (No. 2021YFD1700900) and the Science and Technology Key Project of Henan (No. 232102110064).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Temperature and precipitation during wheat-growing period in 2019–2020 (A) and 2020–2021 (B).
Figure 1. Temperature and precipitation during wheat-growing period in 2019–2020 (A) and 2020–2021 (B).
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Figure 2. Effects of nitrogen application rate on the number and area of vascular bundles and mechanical tissue thickness of the stems of two types of gluten wheat (2020–2021). Small vascular bundle number (A), large vascular bundle number (B), small vascular bundle area (C), large vascular bundle area (D), and mechanical tissue thickness (E). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
Figure 2. Effects of nitrogen application rate on the number and area of vascular bundles and mechanical tissue thickness of the stems of two types of gluten wheat (2020–2021). Small vascular bundle number (A), large vascular bundle number (B), small vascular bundle area (C), large vascular bundle area (D), and mechanical tissue thickness (E). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
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Figure 3. Effects of nitrogen application rate on stem’s mineral nutrients in two types of gluten wheat (2020–2021): K (A), Ca (B), Mg (C), and Fe (D). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
Figure 3. Effects of nitrogen application rate on stem’s mineral nutrients in two types of gluten wheat (2020–2021): K (A), Ca (B), Mg (C), and Fe (D). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
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Figure 4. Effects of nitrogen application rate on the yield and components of two types of gluten wheat in 2019–2020 (A) and 2020–2021 (B). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
Figure 4. Effects of nitrogen application rate on the yield and components of two types of gluten wheat in 2019–2020 (A) and 2020–2021 (B). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
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Figure 5. Effects of nitrogen application rate on grain quality of two types of gluten wheat (2020–2021). Crude protein (A), wet gluten (B), starch (C), sedimentation value (D), flour extraction (E), and bulk density (F). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
Figure 5. Effects of nitrogen application rate on grain quality of two types of gluten wheat (2020–2021). Crude protein (A), wet gluten (B), starch (C), sedimentation value (D), flour extraction (E), and bulk density (F). Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05). p < 0.01: The interaction between nitrogen levels and varieties.
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Figure 6. Structural equation modeling of the effects of nitrogen application on yield, quality, and flexural resistance in medium-gluten wheat. *, significant at p < 0.05; **, significant at p < 0.01.
Figure 6. Structural equation modeling of the effects of nitrogen application on yield, quality, and flexural resistance in medium-gluten wheat. *, significant at p < 0.05; **, significant at p < 0.01.
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Figure 7. Structural equation modeling of the effects of nitrogen application on yield, quality, and flexural resistance in strong-gluten wheat. *, significant at p < 0.05; **, significant at p < 0.01.
Figure 7. Structural equation modeling of the effects of nitrogen application on yield, quality, and flexural resistance in strong-gluten wheat. *, significant at p < 0.05; **, significant at p < 0.01.
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Table 1. Lodging resistance index and its related parameters for four wheat varieties at the fruit development stage under different nitrogen rates from 2019 to 2020.
Table 1. Lodging resistance index and its related parameters for four wheat varieties at the fruit development stage under different nitrogen rates from 2019 to 2020.
VarietyN Rate (kg·ha−1)Plant Height (cm)Stem Diameter (mm)Gravity Center Height (cm)Stem Breaking Strength (N)Lodging Resistance Index
YM49-198N062.9 d4.8 a37.2 d13.1 a35.1 c
N12069.3 c4.7 a40.6 c12.7 ab31.1 b
N18072.4 bc4.7 a44.7 ab12.8 ab 28.5 ab
N24075.8 ab4.7 a43.8 b9.3 b21.2 a
N36077.2 a3.9 b47.2 a4.5 c9.5 a
Mean71.5 BC4.5 AB42.7 B10.5 B25.1 B
JM325N062.1 d4.5 a36.0 c20.1 a55.9 a
N12073.7 c5 a42.8 b11.9 b27.6 b
N18080.1 b4.8 a48.1 a13.4 ab27.6 b
N24085.3 a4.7 a47.8 a10.9 b22.8 b
N36081.1 ab4.1 a47.8 a11.6 b24.4 b
Mean76.5 A4.6 AB44.5 A13.6 A31.6 A
XN979N057.4 c4.9 a32.7 c9.9 a30.4 a
N12068.7 b5.2 a42.4 b9.4 a22.2 b
N18073.7 ab4.7 a45.6 b9.2 a 20.3 b
N24072.9 ab4.6 a49.4 a6.4 b13.0 c
N36075.6 a5.1 a45.3 b5.6 b12.3 c
Mean69.6 C4.9 A43.1 B8.1 C19.6 C
JM44N064.4 b4.6 a36.4 d12.8 a35.0 a
N12068.8 b4.3 a40.7 c12.2 ab 30.0 ab
N18074.6 a4.6 a40.3 c10.3 bc25.6 b
N24077.2 a3.9 a47.7 a8.3 cd17.6 c
N36076.5 a4.1 a44.0 b7.3 d16.6 c
Mean72.3 B4.3 B41.8 B10.2 B25 B
N 17.8 **3.7 *5.3 **15.5 **16.0 **
V 79.0 **ns74.6 **17.0 **44.5 **
N × V 2.1 **ns4.0 **ns3.5 **
*, significant at p < 0.05; **, significant at p < 0.01; ns, nonsignificant at the p > 0.05 level. Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05).
Table 2. Lodging resistance index and its related parameters for four wheat varieties at the fruit development stage under different nitrogen rates from 2020 to 2021.
Table 2. Lodging resistance index and its related parameters for four wheat varieties at the fruit development stage under different nitrogen rates from 2020 to 2021.
VarietyN Rate (kg·ha−1)Plant Height (cm)Stem Diameter (mm)Gravity Center Height (cm)Stem Breaking Strength (N)Lodging Resistance Index
YM49-198N074.2 b4.3 a42.9 b13.9 a32.5 a
N12076.3 b3.9 a44.8 ab10.6 b23.7 b
N18078.8 a4.2 a47.7 a6.2 c12.9 c
N24075.8 b4.2 a45.3 ab5.5 cd12.2 cd
N36077.7 a4.2 a46.0 ab4.5 d9.7 d
Mean76.5 B4.2 A45.3 D8.1 A18.2 A
JM325N077.6 b4.3 a44.8 c13.2 a29.5 a
N12087.5 a4.1 a47.8 b9.7 b20.2 b
N18088.5 a4.2 a51.4 a6.9 c13.4 c
N24086.8 a4.3 a50.5 a7.0 c13.9 c
N36090.6 a4.1 a50.7 a5.6 c11 c
Mean86.2 A4.2 A49.0 B8.5 A17.6 A
XN979N067.2 c3.9 a38.7 d8.6 a22.1 a
N12078.1 b4.4 a47.2 c6.8 b14.3 b
N18077.2 b4.0 a45.1 b5.9 b13 b
N24079.0 b4.0 a50.3 a4.0 c7.9 c
N36082.1 a4.3 a51.8 a3.7 c7.2 c
Mean76.7 B4.1 A46.6 C5.8 C12.9 C
JM44N070.2 c3.1 b41.0 d10.8 a26.3 a
N12087.7 b4.4 a47.8 c7.5 b15.8 b
N18089.8 b4.0 a50.3 c6.5 bc12.9 c
N24087.3 b3.9 a53.7 b6.3 bc11.7 cd
N36093.0 a4.2 a59.0 a5.9 c10 d
Mean85.6 A3.9 A50.4 A7.4 B15.3 B
N 147.6 **ns39.7 **33.8 **25.0 **
V 112.0 **ns89.0 **139.0 **192.7 **
N × V 10.4 **3.1 **12.2 **5.7 **3.9 **
**, significant at p < 0.01; ns, nonsignificant at the p > 0.05 level. Uppercase letters represent significant differences between varieties; lowercase letters represent significant differences between treatments (LSD test, p < 0.05).
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Hu, X.; Tian, P.; Fu, W.; Tian, Z.; Du, M.; Chang, Z.; Ye, Y.; Meng, X.; Wang, Y. Effects of Nitrogen Fertilizer Application on the Lodging Resistance Traits, Yield, and Quality of Two Gluten Types of Wheat. Agriculture 2025, 15, 637. https://doi.org/10.3390/agriculture15060637

AMA Style

Hu X, Tian P, Fu W, Tian Z, Du M, Chang Z, Ye Y, Meng X, Wang Y. Effects of Nitrogen Fertilizer Application on the Lodging Resistance Traits, Yield, and Quality of Two Gluten Types of Wheat. Agriculture. 2025; 15(6):637. https://doi.org/10.3390/agriculture15060637

Chicago/Turabian Style

Hu, Xueling, Peiyu Tian, Wen Fu, Zhihao Tian, Mengdi Du, Zhishang Chang, Youliang Ye, Xiangping Meng, and Yang Wang. 2025. "Effects of Nitrogen Fertilizer Application on the Lodging Resistance Traits, Yield, and Quality of Two Gluten Types of Wheat" Agriculture 15, no. 6: 637. https://doi.org/10.3390/agriculture15060637

APA Style

Hu, X., Tian, P., Fu, W., Tian, Z., Du, M., Chang, Z., Ye, Y., Meng, X., & Wang, Y. (2025). Effects of Nitrogen Fertilizer Application on the Lodging Resistance Traits, Yield, and Quality of Two Gluten Types of Wheat. Agriculture, 15(6), 637. https://doi.org/10.3390/agriculture15060637

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