Micro-sprinkler irrigation with optimal nitrogen application enhances winter wheat yield by regulating flag leaf physiology and root growth in the Huaibei Plain

Limeng Zhang; Huaiwei Li; Bowen Li; Zeen Li; Zifan Hu; Siqi Wang; Chunsheng Yao; Yinghua Zhang; Zhimin Wang; Jincai Li; He Song; Jinpeng Li; Limeng Zhang; Huaiwei Li; Bowen Li; Zeen Li; Zifan Hu; Siqi Wang; Chunsheng Yao; Yinghua Zhang; Zhimin Wang; Jincai Li; He Song; Jinpeng Li

doi:10.48130/tia-0025-0014

A two-year field experiment was conducted to explore the effects of conventional flood irrigation (CI), and micro-sprinkler irrigation (SI) under three topdressing nitrogen levels (N1, 45 kg·ha⁻¹; N2, 90 kg·ha⁻¹; N3, 135 kg·ha⁻¹) on flag leaf physiology, root growth, and grain yield (GY) of winter wheat in the Huaibei Plain (HP). Results showed that higher N significantly decreased the soil moisture content, but SI had higher moisture than CI at the same N level. N2 and N3 significantly increased leaf area index (LAI) compared with N1, and the LAI in SI was higher than in CI. Chlorophyll content in N2 and N3 was significantly higher than in N1 during the grain filling stage, and SI further enhanced the content under the same N level—likely because higher N increased flag leaf antioxidant enzyme activity, while SI outperformed CI. At the anthesis stage, N2 had the highest root length density below 30 cm soil depth, with SI exceeding CI at the same N level. Therefore, N2 and N3 significantly increased dry matter accumulation at maturity (DMM), and the contribution rate of post-anthesis dry matter accumulation to GY (CR). SI further enhanced DMM and CR vs CI at the same N level. Compared to N1, N2 and N3 increased GY by 23.52% and 23.13%, respectively. SI significantly enhanced GY, ranging from 6.10% to 16.99% than CI. Overall, micro-sprinkler irrigation with optimal nitrogen application at jointing and booting stages effectively promoted root growth, enhanced leaf stress resistance, and delayed leaf senescence during grain filling, resulting in improved wheat yield in the HP.

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Introduction

Wheat, a staple global food crop, holds significant economic and nutritional importance, and plays a vital role in worldwide food production and supply^[1,2]. The Huaibei Plain (HP), a primary wheat-growing region in China, lies at the southernmost part of the Huang-Huai-Hai Plain. With ample light and favorable temperature, the HP is suitable for wheat cultivation, and has an annual wheat-planting area of around 2.2 × 10⁵ hectares. Securing high and stable wheat yields in this region is of great significance for ensuring national food security. However, the HP's dominant lime concretion black soil hardens and cracks under severe water deficiency, while excessive moisture from over-irrigation or heavy rainfall impairs its aeration. Regardless, this soil's inherent properties drive rapid soil moisture evaporation. These issues threaten high and stable wheat yields in the region^[3]. Meanwhile, uneven precipitation in the growing season often leads to periodic droughts at wheat's critical growth stages, thereby limiting its yield potential^[4]. Particularly during the jointing to booting stage, wheat frequently experiences insufficient soil moisture or even drought stress. In addition, nitrogen (N) is the most important nutrient element affecting wheat growth and yield formation^[5]. Currently, in the HP, the prevalent practices in wheat production of farmers involves conventional flood irrigation and over-application of nitrogen, leading to wastage of both water and nitrogen resource, and even hindering high wheat yields^[6]. Therefore, exploring appropriate water-nitrogen management techniques is essential for achieving synergistic improvement of water and nitrogen use efficiencies, and the coordinated enhancement of wheat yield in the HP.

As one of the crops with high water requirements, wheat needs over 400 mm of total water consumption for high yields^[7]. Proper irrigation can promote wheat root growth, significantly enhance the uptake and utilization of soil water, increase dry matter accumulation in the population, and ultimately lead to higher grain yield^[8,9]. Studies have shown a close relationship between wheat yield and post-anthesis dry matter accumulation from leaf assimilates. Maintaining optimal soil moisture during critical growth stages delays post-anthesis leaf senescence, significantly enhances photosynthetic capacity, promotes the accumulation and translocation of photosynthetic products to grains, and thereby improves wheat grain weight and yield^[7]. However, soil water deficit can disrupt reactive oxygen species (ROS) metabolism in plants, leading to excessive malondialdehyde (MDA) accumulation, impaired cell structure and function, and rapid degradation of leaf chlorophyll content^[10]. These changes reduce photosynthetic capacity and decrease dry matter accumulation^[11]. To mitigate the adverse effects of external stress, plants rapidly synthesize antioxidant enzymes when stress occurs, such as superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), reducing MDA accumulation^[12]. As the primary organs for water and nutrient acquisition, roots play a vital role in wheat productivity. A well-developed root system ensures a sustained supply of water and nutrients during the post-anthesis period, facilitating assimilate partitioning to grains—a pivotal mechanism underpinning high grain yield. Proper irrigation regimes can promote wheat root growth and deep penetration, thereby enhancing nutrient uptake and efficient utilization^[13]. As a critical period for wheat root development, irrigation during the jointing and booting stages can significantly promote root growth, enhance SOD and POD activity in post-anthesis flag leaf, reduce MDA content, delay leaf senescence, and ultimately improve wheat yield^[14]. Consequently, adopting an appropriate irrigation regime is crucial for promoting wheat root growth, enhancing leaf productivity, and increasing dry matter accumulation, thereby improving yield.

Nitrogen plays a vital role in crop growth and development. In the Huang-Huai-Hai Plain region, wheat is typically supplied with nitrogen application ranging from 180 to 240 kg·ha⁻¹ under conventional flood irrigation, with split between pre-sowing basal fertilization and jointing-stage topdressing^[15]. Research has confirmed that nitrogen application significantly affects wheat growth and development, and optimized nitrogen application can increase effective tillers, kernels per spike, and grain weight, thereby improving crop yield^[16−18]. Excessive nitrogen application, however, may reduce crop yield; additionally, it increases residual nitrate nitrogen in the soil profile, leading to nitrogen leaching, groundwater pollution, and potential harm to the local ecological environment^[19]. Proper nitrogen application primarily enhances crop photosynthetic capacity, delays plant senescence, and promotes the accumulation and translocation of assimilates, thus improving yields^[20]. Conversely, either insufficient or excessive nitrogen application can inhibit root growth, increase lodging risk, delay maturity, reduce photosynthate accumulation, and lower yields^[21]. Micro-sprinkler irrigation, a widely adopted technique in wheat production, enables frequent, low-volume water and fertilizer supply tailored to crop water and nutrient demands. Research has shown that, compared to conventional flood irrigation practices, the integration of water and N-fertilizer under micro-sprinkler irrigation, significantly increases wheat yields by increasing the leaf area index (LAI), maintaining high flag leaf chlorophyll content, and improving photosynthetic performance^[22]. However, the effects of nitrogen application under micro-sprinkler irrigation on dry matter production and yield formation of winter wheat in the lime concretion black soil region of the HP region remain unreported.

Research indicates that the interaction between irrigation and nitrogen application, particularly the amount and timing of topdressing, significantly influences wheat yields^[22,23]. An appropriate increase in topdressing nitrogen application can promote the ability of wheat roots to utilize deep soil water, which in turn effectively improves wheat drought resistance, promotes wheat growth and dry matter accumulation, and mitigates the adverse impacts of water deficit on wheat yields^[24]. Micro-sprinkler irrigation technology ensures adequate soil moisture and nitrogen availability during critical growth stages, reduces nutrient leaching, promotes root development, and enhances wheat's absorption and utilization of soil water and nutrients^[25]. Moreover, timely and adequate irrigation and topdressing nitrogen application can increase flag leaf chlorophyll content and LAI after anthesis, promoting dry matter accumulation, and significantly boosting wheat yields^[7,22]. To summarize, the following research hypothesis is proposed: in the HP region, micro-sprinkler irrigation with optimized nitrogen application ensures a suitable water supply during key wheat growth stages, enhances the stress tolerance of post-anthesis leaf, delays the senescence both leaf and root, and thereby promotes dry matter accumulation of wheat, ultimately increasing grain yield. To verify this hypothesis, a field experiment was conducted during the 2022–2023 and 2023–2024 wheat growing seasons. The experimental design comprised two irrigation methods (conventional flood irrigation and micro-sprinkler irrigation) combined with three topdressing nitrogen levels (45, 90, and 135 kg·ha⁻¹) under a basal nitrogen application of 112.5 kg·ha⁻¹ for each irrigation method. This study examined the effects of different nitrogen application levels under the two irrigation methods on: (1) the spatiotemporal dynamics of soil moisture during the growing season; (2) the physiological traits associated with flag leaf senescence and root distribution; and (3) dry matter accumulation and grain yield of winter wheat. The findings of this study are expected to provide theoretical and technical support for achieving stress-resilient, high-yield wheat production in the HP region.

Discussion

Effects of micro-sprinkler irrigation combined with appropriate nitrogen application level on leaf physiological and dry matter accumulation of wheat

Leaves, as the primary photosynthetic organs in crops, exhibit a direct relationship between their photosynthetic capacity and key physiological parameters including leaf area index (LAI), chlorophyll content, and antioxidant capacity^[32]. The present findings confirm this (Fig. 10). Research has shown that supplementary irrigation and topdressing nitrogen application during the key growth stages can significantly increase antioxidant enzyme activity in flag leaf, maintain high chlorophyll content after anthesis, delay leaf senescence, and promote dry matter accumulation of wheat^[10]. In this study, micro-sprinkler irrigation, combined with optimal topdressing nitrogen application (SIN2) significantly increased LAI at anthesis (Fig. 4), enhanced antioxidant enzyme activity at the anthesis and grain filling stages (Fig. 6), reduced MDA content (Fig. 7), significantly increased flag leaf chlorophyll content (Fig. 5), and ultimately increased wheat post-anthesis dry matter accumulation (Table 2). This is consistent with a previous report that rational water and nitrogen management can extend the functional period of leaves, enhance crop photosynthetic capacity, and thus promote grain yield (GY)^[33].

An optimal LAI serves as a critical regulator of crop source-sink relationships. Previous research has demonstrated that the elevated topdressing nitrogen rates promote vegetative organ development and leaf expansion, thereby enhancing LAI^[34]. In this study, under the same irrigation method, the N2 significantly increased LAI of the anthesis stage compared with N1 (Fig. 4), but there was no significant difference between N2 and N3. Under the same topdressing nitrogen level, micro-sprinkler irrigation significantly increased the LAI compared with conventional flood irrigation (Fig. 4), which is consistent with previous findings^[22,35]. However, the antioxidant enzymes activity under N1 were significantly lower than that under N2 and N3 under the same irrigation method, with no significant difference between N2 and N3 (Fig. 6). This may be because, when leaf nitrogen content is insufficient, chlorophyll degradation causes an imbalance in light energy capture and electron transport, preventing the reactive oxygen species (ROS) from synthesizing sufficient antioxidant enzymes. Conversely, when leaf nitrogen content is excessive, enzyme activity does not further increase due to nitrogen saturation in leaf^[36]. Under identical topdressing nitrogen level, SI significantly enhanced the activities of POD, SOD, and CAT compared to CI (Fig. 6), while significantly reducing MDA content (Fig. 7), which aligns with previous studies^[30]. According to the two-year chlorophyll content data (Fig. 5), at the grain filling stage, the flag leaf chlorophyll content in the 2022–2023 growing season increased significantly with higher nitrogen application levels, with N3 significantly higher than N2. In the 2023–2024 growing season, however, chlorophyll content under N2 and N3 was significantly higher than that under N1, with no significant difference between them. This discrepancy may be because post-anthesis precipitation in 2023–2024 was significantly lower than that in 2022–2023, and temperature was higher than that in 2022–2023 (Fig. 2), resulting in significantly lower soil water at the grain filling stage in the second year (Fig. 3). Under such water-limited conditions, elevated soil nitrogen supply may enhance leaf chlorophyll synthesis^[37,38].

Dry matter accumulation of wheat serves as the foundation for grain yield and is closely linked to crop yield^[39,40]. It is generally acknowledged that approximately 70% of grain yield originates from post-anthesis dry matter accumulation, and increasing the accumulation amount can significantly boost wheat yield^[41]. Rational water and nitrogen management can enhance post-anthesis dry matter accumulation and its contribution to grain yield^[9,22]. In this study, under the same irrigation method, both the DMA and DMM, along with CR, were significantly lower under N1 than under N2 and N3. This phenomenon may be related to lower LAI and decreased leaf stress resistance under low nitrogen conditions, which lead to premature senescence (Figs 4−6). The little difference between N2 and N3 might be attributed to the fact that topdressing nitrogen beyond N2 did not significantly improve leaf material production capacity. However, under the same topdressing nitrogen level, SI significantly increased DMPA and CR compared to CI. This could be related to the significant increase in post-anthesis SWC (Fig. 3), flag leaf chlorophyll content (Fig. 5), and antioxidase enzyme activities under SI (Fig. 6).

Effects of micro-sprinkler irrigation combined with an appropriate nitrogen application level on wheat root distribution
Roots are the main organs for crops to absorb water and nutrients from soil, and their growth and development directly influence above-ground plant growth. In this study, it was also found that the root growth is closely related to wheat yield (Fig. 9). Current research indicates that the rational irrigation regimes and N-fertilizer application can regulate root growth, facilitating a reasonable distribution of roots in the soil profile^[42−44]. Concurrently, an optimal topdressing nitrogen level can significantly activate nitrogen signaling pathways, and induce roots to penetrate into the deep soil layer^[45]. In this study, compared with CI, SI significantly promoted the root growth and deep penetration under the same nitrogen level, with significantly higher root length density (RLD) in surface and deep layers at the anthesis stage (Fig. 8). This finding differs from previous studies, potentially due to soil type effects^[29]. In the HP region, the soil is mainly lime concretion black soil with clayey characteristics. The conventional flood irrigation at the jointing stage may reduce soil permeability, thus affecting root growth, while micro-sprinkler irrigation with split and small irrigation volumes enables uniform water infiltration, maintains soil permeability, and creates a more favorable environment for root development. Deep roots can not only enhance water absorption capacity in the later period but also delay leaf senescence by transporting cytokinins upward^[46]. In this study, under the same irrigation method, N2 treatment had the most surface roots and promoted deep root distribution, which is consistent with previous studies^[29,47]. In addition, some studies have shown that root growth is influenced not only by soil water but also by nitrate nitrogen (NO₃⁻-N) concentration. A low NO₃⁻-N concentration can significantly increase auxin content in roots and enhance root elongation^[48]. However, an excessively low nitrogen application level may restrict root growth and development, inhibit root penetration into deep layers, and reduce the ability of the crop to obtain water and nutrients from the soil. In summary, micro-sprinkler irrigation combined with an appropriate nitrogen application (SN2) may optimize root architecture by improving the root zone environment. Simultaneously, the coordinated development of roots and above-ground parts promote dry matter accumulation; root penetration to deep layers enhances the absorption capacity of plants for water and nutrients, thereby supporting continuous photosynthesis of post-anthesis flag leaf, facilitating the translocation of assimilates to grains, and ultimately increasing the CR (Table 2). Furthermore, micro-sprinkler irrigation combined with optimal nitrogen application may regulate root growth by changing soil NO₃⁻-N concentration during the key period of root growth (from jointing to anthesis). However, future studies need to further investigate whether NO₃⁻-N concentration plays an important role in promoting root growth and development under optimal nitrogen application level for micro-sprinkler irrigation, thereby contributing to yield improvement.

Effects of micro-sprinkler irrigation combined with appropriate nitrogen application level on wheat grain yield (GY) and its components
GY of wheat is determined by three components: spike number (SN), kernels per spike (KN), and thousand-grain weight (TGW), all closely associated with water and nitrogen supply^[49]. Achieving high GY typically requires increasing KN and TGW while maintaining adequate SN^[25]. The jointing and booting stages are key periods for wheat KN formation, and represent sensitive periods for water and nutrient availability. Rational water and N-fertilizer application during this period can promote root penetration, thereby enhancing the material production capacity of wheat, which positively influences increases in KN and GY^[10]. Research has shown that SI can effectively extend the post-anthesis leaf photosynthetic duration through multiple small-volume water applications, improve the post-anthesis dry matter accumulation and CR, and ultimately increase TGW and GY^[21]. In this study, under the same irrigation method, the GY in N1 was significantly lower than that in N2 and N3, with no significant difference between N2 and N3 (Table 3). This is because insufficient nitrogen application may reduce wheat spike differentiation and pollen fertility, thereby affecting SN and KN^[50]. The present study indicated that the SN in SI was significantly higher than CI under N2 during the 2022–2023 growing season of winter wheat, this may be due to the lower rainfall from the jointing to booting period in this year (Fig. 2). However, previous research result indicated that beyond a certain nitrogen application level, further increasing nitrogen application does not necessarily enhance yield^[51]. This experiment proved that micro-sprinkler irrigation with integrated water and N-fertilizer combined with an appropriate nitrogen application level, extends the grain formation period through water and nitrogen synergy, ensures water supply during post-anthesis grain filling via split irrigation, increases the TGW, and ultimately promotes GY. Additionally, the lime concretion black soil area in the HP has the characteristics of heavy soil texture in rainy seasons and easy hardening in dry seasons. Under the micro-sprinkler irrigation with integrated water and N-fertilizer combined with an appropriate nitrogen application level (SIN2) provides sufficient nutrients at the key stages for wheat, maintains high soil water content (Fig. 3), promotes root growth into deep layers (Fig. 8), and achieves root-shoot coordination. This not only improves pre-anthesis photosynthetic production capacity and increases the LAI at anthesis (Fig. 4) but also delays post-anthesis leaf senescence (Figs 5−7) and promotes dry matter accumulation and translocation (Tables 2, 3), thereby significantly increasing wheat KN and TGW, and ultimately boosting GY (Table 3).

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Treatment	Irrigation amount (mm)			N application amount (kg·ha⁻¹)
Treatment	Jointing stage	Booting stage	Total	Base fertilizer	Jointing stage	Booting stage	Total
CIN1	60.0	/	60.0	112.5	45.0	/	157.5
CIN2	60.0	/	60.0	112.5	90.0	/	202.5
CIN3	60.0	/	60.0	112.5	135.0	/	247.5
SIN1	30.0	30.0	60.0	112.5	22.5	22.5	157.5
SIN2	30.0	30.0	60.0	112.5	45.0	45.0	202.5
SIN3	30.0	30.0	60.0	112.5	67.5	67.5	247.5

Treatment	Dry matter accumulation in 2022–2023				Dry matter accumulation in 2023–2024
Treatment	DMA (kg·ha⁻¹)	DMM (kg·ha⁻¹)	DMPA (kg·ha⁻¹)	CR (%)	DMA (kg·ha⁻¹)	DMM (kg·ha⁻¹)	DMPA (kg·ha⁻¹)	CR (%)
CIN1	11,646.52b	16,974.12d	5,327.60f	71.30c	11,533.30b	16,081.33c	4,548.03c	64.80c
CIN2	11,944.90ab	18,582.00b	6,637.10c	75.51b	12,452.80a	18,503.30b	6,050.50b	67.69b
CIN3	12,004.53ab	18,058.93bc	6,054.40d	72.60c	12,851.17a	18,384.93b	5,533.77b	66.38b
SIN1	11,689.62b	17,475.42cd	5,785.80e	74.12bc	10,431.20c	16,404.80c	5,973.60c	68.11c
SIN2	12,169.93a	20,067.83a	7,897.90a	78.83a	11,764.13b	19,185.27a	7,421.13a	74.54a
SIN3	12,219.67a	19,513.67a	7,294.00b	75.82b	11,829.00b	18,971.77a	7,142.77a	76.67a
ANOVA
I	ns	**	**	**	**	**	**	**
N	**	**	**	**	**	**	**	**
I×N	ns	ns	**	ns	ns	ns	ns	*
Note: CI, conventional flood irrigation; SI, micro-sprinkler irrigation; N1, N2, and N3 represent topdressing nitrogen with 45 kg·ha⁻¹, 90 kg·ha⁻¹, and 135 kg·ha⁻¹ respectively. DMA, dry matter accumulation at anthesis stage. DMM, dry matter accumulation at maturity stage; DMPA, post-anthesis dry matter accumulation; CR, the contribution rate of dry matter accumulation post-anthesis to grain yield. Different lowercase letters within a column indicate significant differences at p ≤ 0.05. I indicates the irrigation methods, N indicates the nitrogen application levels, and I × N indicates the interaction. * and ** represent the significant difference at the 0.05 and 0.01 levels, respectively, ns represents that the difference is not significant.

Treatment	2022–2023				2023–2024
Treatment	GY (kg·ha⁻¹)	SN (× 10⁴·ha⁻¹)	KN	TGW (g)	GY (kg·ha⁻¹)	SN (× 10⁴·ha⁻¹)	KN	TGW (g)
CIN1	7,517.78 d	573.48 c	37.19 d	42.18 d	7,019.30 d	562.55 b	36.88 d	42.60 d
CIN2	8,790.80 b	609.18 b	40.55 b	45.21 b	8,319.44 b	604.03 a	40.19 b	44.82 b
CIN3	8,714.15 b	615.85 ab	40.43 b	45.17 b	8,321.11 b	602.70 a	40.15 b	44.31 b
SIN1	7,861.71 c	579.68 c	39.19 c	43.58 c	7,623.81 c	564.22 b	38.51 c	43.48 c
SIN2	10,022.73 a	625.88 a	43.48 a	47.72 a	9,940.51 a	607.11 a	41.34 a	47.49 a
SIN3	9,958.53 a	627.19 a	42.91 a	47.19 a	9,960.48 a	605.22 a	41.16 a	47.20 a
ANOVA
I	***	**	***	***	***	ns	***	***
N	***	***	***	***	***	***	***	***
I × N	*	ns	ns	ns	***	ns	ns	***
Note: CI, conventional flood irrigation; SI, micro-sprinkler irrigation; N1, N2 and N3 represent topdressing nitrogen with 45 kg·ha⁻¹, 90 kg·ha⁻¹, and 135 kg·ha⁻¹ respectively. GY, grain yield; SN, spike number; KN, kernel number per spike; TGW, thousand-grain weight. I indicates the irrigation methods, N indicates the nitrogen application levels, and I × N indicates the interaction. Different lowercase letters within a column indicate significant differences p ≤ 0.05 , , and ** indicate significant difference at p ≤ 0.05, ≤ 0.01, and ≤ 0.001, respectively, ns represents that the difference is not significant.

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Micro-sprinkler irrigation with optimal nitrogen application enhances winter wheat yield by regulating flag leaf physiology and root growth in the Huaibei Plain

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