1. Introduction
Rice (
Oryza sativa L.) is one of the most important staple crops [
1,
2] feeding approximately three billion people in the world [
3]. Faced with an increasing demand for rice, improving rice yield potential has been regarded as the primary objective of rice breeding in many countries for several decades [
4,
5]. In 1996, a “super” rice program was launched to breed rice varieties with high yield potential in China [
6]. As of 2019, 132 “super” rice varieties were released by the Ministry of Agriculture of China (
http://www.ricedata.cn/variety/superice.htm). These “super” rice cultivars, especially the japonica/indica hybrid rice, have made tremendous contributions to food security in China. However, the promotion of japonica/indica hybrids for large-scale applications has been greatly limited by various factors, including the low rate of grain filling [
7] and sterility in the first generation [
8]. In recent years, great progress has been achieved in solving some of these obstacles, and high-yielding japonica/indica hybrids have been bred successfully and made available in China [
9]. The Yongyou japonica/indica hybrid series, which is a late-maturity type, was one of the representative high-yielding japonica/indica hybrids. For example, Yongyou 12 has achieved 15 t ha
−1 yield performance in production for two executive years in Zhejiang Province, China [
10,
11].
Nitrogen (N), the most critical nutrient for rice growth, is required in larger amounts than other nutrients [
12]. To improve yield performance, most Chinese farmers apply N fertilizer in amounts that exceed the demand for rice growth [
13]. However, the N recovery efficiency (NRE) of rice in paddy soils of southern China is generally low, ranging from 20% to 40% [
14]. Lodging and yield losses can be caused by luxury absorption due to the overapplication of N fertilizer [
15,
16]. In addition, the excessive use of N fertilizer also enhances N losses through runoff, leaching, and volatilization, into the environment, which has a negative effect on ground water, surface water, and the atmosphere [
17,
18]. Several studies have shown that an optimum amount of nitrogen application is essential for high rice yield [
5,
19,
20,
21]. For example, Zhu et al. [
5] found that the optimum amount of nitrogen fertilizer for increasing grain yields of japonica rice cultivars Nanjing 9108 and Nanjing 5055 was 262~300 kg ha
−1. To date, however, little information is available on the yield of japonica/indica hybrid rice in response to different nitrogen levels. More information is needed to find the suitable nitrogen application rate to obtain optimal yield of japonica/indica hybrid rice.
The Yongyou japonica/indica hybrid rice is highly heterotic and superior to japonica conventional rice in China, in terms of grain yield, biomass production, and nitrogen accumulation. However, this superiority was observed at a single N application rate (262.5 kg ha
−1) [
22,
23]. Whether it also exists at a higher or lower N application rates under the same field conditions remains unclear. Hence, in this study, we assess the effects of different nitrogen application rates on the agronomic performance of japonica/indica hybrid cultivars, including yield, yield component, dry matter weight, and N uptake as compared with japonica cultivars at various growth stages.
4. Discussion
Nitrogen application is one of the most important crop management practices for achieving a higher grain yield [
25]. Several studies have focused on the effects of the N rates on the grain yield of rice [
5,
19,
26]. In this study, higher N application rates resulted in higher grain yields of Yongyou 12, Yongyou 538, and Xiushui 134 in 2016, 2017, and 2018, except for Yongyou 538 in 2018 (
Table 4). However, no significant increase was observed with the increasing N application rates. According to Chen et al. [
27] and Li et al. [
28], remobilization of N from vegetative tissue to grain could significantly contribute to crop yield. No significant difference among the relative high rate treatments was mainly attributed to the less efficient N remobilization at high N rates. By contrast, a slight decrease in grain yield of Jia 58 was observed when the N rates increased from 300 to 375 kg ha
−1 (
Table 4). Similar results have also been observed in previous studies [
5,
19], revealing that an excessive N application rate has no contribution to the achievement of high grain yield. However, no significant difference was observed between N225 and N300 treatments. Furthermore, nitrogen reduction is recommended due to environmental pollution problems. Therefore, a N application rate of 225 kg ha
−1 is essential for Jia 58.
Significantly higher yield potential has been found in japonica/indica hybrid rice than in inbred rice [
22,
23,
29]. Likewise, in this study, Yongyou 12 and Yongyou 538 had a significant yield advantage over Xiushui 134 and Jia 58 for the five N treatments (
Table 4), which mainly resulted from the heterosis and the longer total growth duration (
Table 2). On average, the grain yield of japonica/indica hybrid rice was higher than that of japonica rice by 75.6% at N0, 57.2% at N150, 41.1% at N225, 38.3% at N300, and 45.8% at N375, across the three planting years. These findings suggest application of N fertilizer would not facilitate to realize a higher grain yield for japonica/indica hybrid rice as compared with the japonica rice. However, this difference in grain yield was not affected by N supplies and environmental conditions, suggesting that grain yield was highly genetically controlled. Thus, in practice, a farmer could simply choose high grain yield varieties such as Yongyou 12 or Yongyou 538 to achieve high grain yield, without much concern for the environmental conditions. However, higher yields of Yongyou 12 and Yongyou 538 were primarily achieved when high N was supplied. Therefore, farmers need to optimize the N management to achieve both high grain yield and less nitrogen input. Interestingly, except for Xiushui 134, the grain yields of the four rice varieties decreased in the three study years, especially for Yongyou 12 and Yongyou 538 (
Table 2 and
Figure 2), which could be due to the appearance of Ustilaginoidea virens in 2017 and 2018.
According to Liu et al. [
30], higher total spikelets number, which led to a larger sink capacity and large panicle size, is beneficial for the increase of yield potential. In this study, we observed significant higher spikelet per panicle in Yongyou 12 and Yongyou 538 as compared with Xiushui 134 and Jia 58 (
Table 5), which contributed to the higher grain yields of Yongyou 12 and Yongyou 538, in terms of yield components, as suggested by Meng et al. [
9]. However, this was contradictory to the previous studies that reported the hybrid rice tended to have higher grain filling percentage and grain weight than the inbred rice [
31,
32]. The discrepant results of these studies could be due to the different types of rice cultivars.
To obtain high grain yields, the appropriate proportion of dry matter accumulation at each growth stage was important, particularly in the middle and late growth stages [
23]. In this study, dry matter of japonica/indica hybrid rice was higher than that of japonica rice among the five nitrogen application rates at each growth stage (
Figure 3,
Figure 4 and
Figure 5). According to Wei et al. [
22], higher leaf area index and leaf SPAD value of japonica/indica hybrid rice were observed at the heading and maturity stages as compared with japonica rice, suggesting its higher radiation interception. In addition, larger leaf areas increase photosynthetic rates [
33]. This could be the reason that japonica/indica hybrid rice showed higher dry matter accumulation over japonica rice.
High yield productivity of rice is usually accompanied by greater N uptake [
34]. In this study, we observed that as compared with Xiushui 134 and Jia 58, Yongyou 12 and Yongyou 538 accumulated more N at each growth stage (
Figure 6,
Figure 7 and
Figure 8). Similar results were also obtained by Wei et al. [
22] and Wei et al. [
33], who revealed that japonica/indica hybrid rice accumulated more N than japonica conventional rice and indica hybrid rice during the entire growth stages. This phenomenon was mainly attributed to the long duration from jointing to maturity (
Table 2). It is worth mentioned that at the maturity stage, a significant difference was observed between japonica/indica hybrid rice and japonica rice, in terms of SNU, GNU, and TNU. Nutrient absorption in rice is closely related to root characteristics. Rice roots with good morphology and excellent physiology are beneficial to the acquisition of nutrients that maintain crop plants growth [
35,
36]. Several studies have demonstrated that japonica/indica hybrid rice had a stronger and more active root system than japonica conventional rice [
9,
33]. This could explain why japonica/indica hybrid rice absorb more nitrogen than japonica conventional rice. Furthermore, according to the correlation analysis, the combination priority of japonica/indica hybrid rice over japonica conventional rice, in terms of grain yield, N uptake at the jointing stage and the heading stage, in turn, enhanced the absorption of N (
Table 7).
The response to applied N is an important indicator for the evaluation of the N requirements of rice. The response to the N application rate of the four rice varieties was similar, which was that the N uptake significantly increased with increasing N application rates (
Table 6). This was in line with a study that reported the TNU increased in both years as the N application rates increased [
37]. However, nitrogen uptake and N use efficiency varies with different rice cultivars. Several studies have suggested that the hybrid rice has higher N use efficiency than the inbred rice, which primarily ascribed to the higher NHI [
22,
23,
32,
33]. However, in our study, no significant differences in NHI between japonica/indica hybrid rice and japonica rice were observed (
Table 6). In addition, we observed that the NHI of the N375 treatment was relatively low as compared with other treatments, although the difference among the treatments was insignificant, suggesting that a higher N supply can lead to high levels of residual N in straw at maturity. Zhu et al. [
5] reported that AE was decreased with increasing nitrogen levels. Similarly, in the present study, the AE of the four rice varieties decreased with increasing N rates, except for Jia 58 (
Table 6), which was mainly due to low efficiency of the increased nitrogen fertilizer. This result indicated that a high N rate had negative effects on AE.
5. Conclusions
Nitrogen application rates significantly affected the grain yield, dry matter, and N uptake. The highest gain yields of all the rice varieties were obtained under the highest N application rates in the field experiment, except for Jia 58, whose grain yield decreased slightly when the N rates was increased from 300 to 375 kg ha−1, however, no significant difference was observed, and therefore the optimum nitrogen level of Jia 58 was 225 kg ha−1. Further research is needed to assess the effects of nitrogen application rates used in this study, on the grain yield of Yongyou 12, Yongyou 538, and Jia 58. Across N treatments in all planting years, dry matter of Yongyou 12, Yongyou 538, and Xiushui 134 tended to increase with N-fertilizer rates, except for Yongyou 538 in 2018, whereas for Jia 58, it was highest with the N300 treatment. However, across the four rice varieties, N uptake increased significantly with increased N-fertilizer rates at all the growth stages (p < 0.05). In addition, as compared with N0, a decline in AE and NHI was observed with increasing N application rates. We also found that as compared with japonica rice, the japonica/indica hybrid rice had more grain yield, spikelets per panicle, as well as higher dry matter and higher N uptake at all the growth stages, regardless of the N application rate.