1. Introduction
Grasslands provide ecosystem services, including the regulation and storage of water flows, nutrient cycling, and carbon sequestration [
1]. However, the excessive use of the same pasture area with high animal density and without nutrient reposition in the soil has promoted the expansion of degraded pasture areas in many places around the world, which have not recovered naturally [
2]. Brazil maintains the largest commercial cattle herd in the world, with pastures covering approximately 177 million hectares; of this area, 41% shows moderate signs of degradation, while 21% is severely degraded [
3].
From an agricultural perspective, double-cropping systems involving two cash crops in a single agricultural year have gained popularity among farmers due to their potential for short-term profitability [
4]. However, the widespread adoption of soybean and maize monocultures has resulted in low crop diversity and greater uniformity of agricultural landscapes, making cropping systems less efficient and sustainable [
5]. Thus, there is a necessity to adopt recovery practices in grassland and grain cropping areas to decrease or mitigate environmental damage [
6] and promote diversification in production systems so that the cycle of intensification can be broken.
Diversification practices, such as intercropping systems, have been shown to increase the production and stability of cropping systems with reduced environmental impacts. In practice, the intercropping system is the simultaneous growth of two or more crops on the same land [
7]. A strategy developed to increase the diversity of plant species in the grain crop, recover degraded pasture, and produce straw involves the use of tropical grasses during the dry season intercropped with maize [
8].
Brazil’s total maize cultivation spans approximately 22.7 million hectares. The second crop (dry season) accounts for 18.1 million hectares, representing nearly 80% of the country’s total maize output [
9]. Dry season maize intercropped with tropical grass can raise the amount of crop residue and improve nutrient cycling, favoring the soybean crop [
9]. Among the species studied in this intercropping system, the genera
Urochloa and
Megathyrsus showed large amounts of dry biomass [
10]. However, maize grown in the dry season after harvesting the summer crop develops under conditions of high climate variability with restrictions on rainfall, air temperature, and solar radiation [
11]. These restrictions promote reduced soil moisture, which can lead to decreased root expansion and metabolic activity, reducing the capacity of plants to uptake the essential nutrients.
Adopting optimal N management can provide greater soil cover and synchronize the nutrient supply during the period of greatest crop demand in agroecosystems [
12]. Tropical grasses and maize are N-demanding plants, and their low availability in the soil can result in variations in the dry biomass, mineral composition, and grain yield of these plants [
13]. Furthermore, as noted by these authors, crop residue decomposition is slower in the no-tillage system, especially in the high biomass production system.
Low N supply in maize crop stimulates the mineralization of soil organic N and subsequent carbon loss, leading to soil degradation as the soil C/N ratio remains nearly constant. In maize monocultures, a significant portion of N fertilizer is lost to the environment. Although intercropping maize with tropical grasses can reduce N leaching, it may also induce temporary N deficiency [
14]. However, this system promotes long-term nutrient cycling; according to [
15], more than 60% of the nitrogen supplied remains within the system, partitioned between tropical grass biomass, maize residues, and the soil.
While maize–grass intercropping has been proposed to enhance biomass production and nutrient cycling [
6,
7,
8], current evidence is largely derived from systems established under relatively favorable edaphoclimatic conditions [
4,
9]. There remains a critical gap in understanding how nitrogen side-dressing interacts with species-specific competitive dynamics in degraded pasture areas subjected to dry season climatic stress. In such environments, low soil fertility and water limitation may substantially alter nitrogen use and nutrient partitioning patterns [
10,
11,
12]. Although the productivity of
Urochloa and
Megathyrsus species has been documented [
8,
13], direct comparisons of these grasses under identical nitrogen gradients in degraded pasture contexts remain limited. By investigating these interactions across two consecutive dry season crops, this study provides new insights into nitrogen-mediated resilience and nutrient cycling in restoration-oriented production systems.
In this context, we hypothesized that species-specific interactions and N fertilization would result in distinct responses, determining the effectiveness of the intercropping system in mitigating dry-season constraints and enhancing nutrient cycling in degraded pasture areas. Thus, the objective of this study was to evaluate the dry biomass, grain yield, and macronutrient concentrations in maize–tropical grass intercropping as a function of the N rates applied as side-dressing in the dry season in a degraded pasture area.
4. Discussion
When maize was intercropped with Congo grass, the minimum maize heights varied with the N rate applied as side-dressing (
Figure 3a), suggesting that N supply is a primary driver of crop development in intercropping systems [
23]. Conversely, the observation that maize monoculture achieved its highest grain yield without N side-dressing in the first crop can be attributed to the restricted rainfall (178.8 mm;
Figure 1). The uptake of water and N by maize is reduced under low precipitation; consequently, N availability to the plant is compromised, as N moves primarily via mass flow. Under such water-limited conditions, particularly after the V6 stage (six fully expanded leaves), the restriction of N uptake and aboveground biomass production appears to have limited the potential for grain number and mass [
24], rendering additional N fertilization ineffective in that season. Similar to this study, Ref. [
25] observed that when the amount of nitrogen application exceeds a certain value, the maize yield does not increase.
Precipitation in the second crop was higher than in the first crop, which may have influenced the responses of the maize monoculture and maize intercropped with Aruana Guinea grass to the N rate applied as side-dressing, despite the occurrence of frost during the grain-filling period (
Figure 1). In general, maize yields in the current study were low across both growing seasons (
Table 1). This was likely because drought conditions during the late vegetative growth period and the onset of silking (stigma–style exsertion) represent a critical window where limited soil moisture and nutrient availability directly reduce yield potential [
26]. In contrast, Ref. [
27] studying maize cropping system and N rates in summer season, observed higher grain yield (9.2, 8.8 and 9.7 Mg ha
−1) for the maize monoculture, maize intercropped with palisade grass (
Urochloa brizantha cv. Marandu) and maize intercropped with guinea grass (
Megathyrsus maximum cv. Mombaça) at N rates of 113, 156 and 187 kg ha
−1 N respectively. According to Ref. [
28] changes in temperature and precipitation patterns affect biological and enzyme activity rates, which are essential for most soil N transformations. High crop growth rates, yield, and N uptake can be achieved by maintaining optimal soil moisture conditions.
The impact of climatic variables on grain yield in this study is evidenced by the alignment between rainfall patterns and crop performance (
Figure 1 and
Figure 2). When soil moisture becomes a limiting factor, the resulting water deficit restricts nutrient transport and biomass accumulation, ultimately capping the grain yield potential [
29,
30]. Furthermore, the extremely low temperatures recorded during the second crop (
Figure 1) represent a significant thermal stress capable of damaging reproductive structures [
26]. Notably, the presence of Aruana Guinea grass appeared to provide a protective effect for the maize plants during these periods of thermal stress (
Figure 3b). Identifying these environmental interactions within intercropping systems is essential for optimizing nitrogen use efficiency, ensuring a positive nitrogen balance, and developing resilient sustainable production models.
The lowest dry biomass of maize in monoculture occurred at an estimated N rate of 67.11 kg ha
−1 and 156.52 kg ha
−1 (
Figure 3c) at its physiological maturity in the first and second crops indicated that although solar radiation, temperature, and water availability were less favorable for maize growth during this period, its total suppression is not recommended [
31]. According to Ref. [
32], the lack of water reduced the dry biomass in all parts of the maize plant under different N rates. Ref. [
13] observed that during the dry season, maize dry biomass at physiological maturity increased 1.8-fold when intercropped with Congo grass at an N rate of 90 kg ha
−1 compared to the treatment without N. Similarly, the findings of Ref. [
27] revealed that during the summer season, the dry biomass of the maize intercropping system using 200 kg ha
−1 of side-dress N was the highest for the maize–palisade grass intercropping. Moreover, Ref. [
33] emphasized that nitrogen fertilization is crucial for early maize growth, enabling rapid shading of the intercropped grass and minimizing interspecific competition.
The higher dry biomass of Congo grass compared to Aruana Guinea grass at the physiological maturity of maize and at its desiccation in the first crop (
Table 1) suggests differences in the competitive capacity of these grasses for growth factors and interspecific interactions within the rhizosphere [
34]. Similar to the results of this study, Ref. [
9] also observed an increase in the dry biomass production of tropical grasses intercropped with maize in the dry season at the physiological maturity of maize and at its desiccation. These results suggest that the use of these plants could contribute to animal feed and soil protection in the dry season. Ref. [
35] highlighted that Congo grass had the capacity to accumulate a large amount of biomass, maintaining the soil covered in the off-season, which is interesting, as the biomass maintains the soil moisture, enabling soybean growth in succession, especially in crops with prolonged water stress or irregular rainfall distribution.
The lack of response of grass dry biomass in the second crop (
Table 2) can be associated with the stress conditions that occurred during this period. When plants are subjected to stress, their stability is disrupted, including the alteration of metabolic pathways and the production of excess electrons, which increases intracellular levels of reactive oxygen species that damage cellular structures [
36].
The differences in the N supply required to maximize the N concentration in the shoots of maize in monoculture (128.75 kg ha
−1) versus those in maize intercropped with Congo grass (138.10 kg ha
−1) at its physiological maturity in the second crop indicate that it is necessary to adjust N rates applied as side-dressing to meet specific plant demands according to the cropping system (
Figure 4a). According to Ref. [
37] the most N-efficient scenarios are realized when the soil N supply matches the plant’s demand for N throughout the entire growth cycle. Furthermore, Ref. [
31] highlighted possible interspecific competition within the maize–Congo grass intercropping system, which resulted in a reduction in the amount of N taken up by the maize. In this context, Ref. [
15] observed that N management must be tailored to account for these competitive interactions to ensure that grain yield is not compromised.
The P concentration in the maize shoots at physiological maturity in the second crop as a function of the cropping system and N rate applied as side-dressing showed that the plants can use one element to facilitate the acquisition of another (
Figure 4b), suggesting an adequate N rate favored P uptake. P availability may be mediated by rhizosphere interactions, particularly in the intercropping system where the root systems intermingle [
38].
The K and Ca concentrations in the shoots of maize intercropped with Aruana Guinea grass at its physiological maturity in the second crop were favoured by the N supply (
Figure 4c). This could be related to the intermingling of roots from the two intercropped species, which enables the transfer and exchange of substances and signals [
38]. However, as the increase in the N rate promoted an increase in the K concentration in maize shoots, it is necessary to monitor for luxury uptake. However, it is important to note that the observed K concentrations in this study remained within the adequacy ranges for maize, indicating that excessive accumulation did not occur [
21].
The maximization of the Mg concentration in the shoots of maize in monoculture at physiological maturity, in the first and second crops, as a function of the N supply (
Figure 4d) could be associated with the reduction in the dry maize biomass, as the increase in the N supply resulted in a low dry maize biomass. Mg accumulation in plants occurs in greater amounts in the chloroplasts and mitochondria, followed by the cytosol. Furthermore, the vacuole serves as a large Mg pool to maintain the cytosolic Mg balance [
39].
The highest S concentration in the shoots of maize intercropped with Aruana Guinea grass at physiological maturity in the second crop indicated that this grass may have facilitated S absorption by the maize (
Table 4). In addition, this result may be associated with the positive grain yield response of maize to the N rate when intercropped with Aruana Guinea grass (
Figure 3b). S plays an important role in the synthesis of amino acids, proteins, coenzymes, and chlorophyll, consequently changes in its absorption can reduce the grain yield and the concentrations of carbohydrates, starch, and proteins in maize [
40].
The alterations observed in macronutrient concentrations in maize as a function of the cropping system and N supply highlight the need to evaluate maize development under different practices. Furthermore, these variations must be interpreted alongside biomass accumulation, as dilution effects often occur when increased productivity outpaces nutrient uptake [
9].
The N concentration in the shoots of Congo grass intercropped with maize at physiological maturity in the first crop was low when the N supply applied as side-dressing was inadequate (
Figure 5a). Once again, the results reveal that the synchronization of nitrogen supply is essential to ensure a positive nitrogen balance [
37]. However, at the physiological maturity of maize in the second crop, the highest N concentration in the shoots of Aruana Guinea grass and Congo grass intercropped with maize occurred outside the range of rates studied (
Figure 5a). Moreover, Aruana Guinea grass intercropped with maize required 1.45 times more N to reach its maximum N concentration than Congo grass. These responses are important because the N present in the residues of cover crops decreases linearly over time after maize harvest and cover crop management [
41].
The highest P concentration in the shoots of Aruana Guinea grass intercropped with maize at the physiological maturity of maize in the first and second crops (
Table 5 and
Table 6) showed that this grass had a high P absorption capacity. This is particularly significant given that P is an immobile nutrient in the soil and enters the roots primarily through diffusion. Furthermore, these results suggest that Aruana Guinea grass can recover P fractions from the soil even under conditions of low P availability [
42].
The Mg concentration in the shoots of Aruana Guinea grass intercropped with maize in the first and second crops suggests that an inadequate N supply (
Figure 5b), may alter the photosynthetic capacity of this grass, as Mg is a central component of chlorophyll molecules. Similarly, at the physiological maturity of maize, Congo grass shoots exhibited a reduction in the Mg concentration as N supply increased in the second crop (
Figure 5b). The concentrations of Ca and Mg in Aruana guinea as a function of the N rate are likely associated with a dilution effect driven by increased vegetative growth, since Ca is essential for middle lamella and cell wall formation, while Mg is vital for photosynthesis. Furthermore, N supply favored higher S concentrations in Congo grass (
Figure 5c), aligning with the findings of [
9].
The influence of the N rates applied as side-dressing on the N concentration in the shoots of Aruana Guinea grass intercropped with maize at its desiccation in the first crop and that of Congo grass intercropped with maize at its desiccation in the second crop confirms that these intercropping systems are N dependent (
Figure 5a). These results indicate that the effective contribution of N from grasses to the subsequent crop depends on the concentration of this nutrient in the biomass and on the mineralization of the accumulated N [
43]. In addition, Ref. [
44] observed that an intercropping system between monocotyledons could self-regulate soil N levels and optimize nutrient use for succeeding crops. According to Ref. [
45] N primarily exists in highly compacted organic forms, which facilitate slow nutrient release
The P concentration in the shoots of Aruana Guinea grass intercropped with maize at its desiccation in the first and second crops (
Figure 5c) is consistent with the results of [
46], who demonstrated that plants use N to acquire P from organic sources and that N addition could influence phosphatase activity. Ref. [
45] emphasized that P contained in plant residues exists primarily as inorganic P, whereas the organic portion is difficult to decompose.
The N supply increased the K concentration in Congo grass shoots intercropped at its desiccation in the first and second crops (
Figure 5d), indicating enhanced K availability for the subsequent crop. K contained in plant residues exists primarily in an ionic state and easily dissolves easily in water during decomposition process [
45]. Since K in plant residues is released relatively quickly, the adoption of an intercropping system can maintain this nutrient in the soil layers exploited by the roots [
42].
Aruana Guinea grass intercropped with maize showed a greater ability to absorb and return Ca to the system than Congo grass in the first crop, benefiting the subsequent crop (
Table 7). According to Ref. [
41], the highest release of Ca from cover crops intercropped with maize occurs approximately 75 days after maize harvest and cover crop management.
The maximum Mg concentration in Congo grass shoots intercropped with maize at its desiccation in the first and second crops (
Figure 5b) as a function of an adequate N rate suggests that it is necessary to balance N demand and the Mg content to prevent possible deficiencies in this crop. Similarly, Ref. [
13] observed significant effects of N fertilization on the Mg concentration in the Congo grass shoots intercropped with maize.
The macronutrient concentrations in the grass shoots showed that the interaction between the cropping system and the N rate applied as side-dressing is essential for nutrient cycling, particularly for the subsequent crop. This is because the soil remains active throughout the off-season, contributing to system stability [
35]. Regardless of the N rate applied, the macronutrient concentrations in the grass shoots followed the decreasing order: N > K > Mg > Ca > S > P. These results emphasize the importance of grasses intercropped with maize, as the release rates of N, P, K, and Mg from grass residues vary over 120 days after desiccation, with peak release occurring within the first 30 days [
10]. Furthermore, variations in nutrient concentration must be interpreted alongside biomass production to avoid misleading conclusions, as dilution effects were prominent in the high-yielding treatments [
9,
43].