3.1 Results
During a one-year cropping cycle, maize grew by uptaking the mineral N that is mineralized from the soil organic matter. The maize biomass increases with the absorption rate and is maximum for intermediate preferences for NO3− vs. NH4+ that are higher than 0.5 (preference for NH4+) (Fig. 2). The increase in maize biomass with the uptake rate increases steeply for low uptake rate values and then reaches a plateau. This general pattern is the same for the three mineralization rates (Fig. 2), but the maximum maize biomass increases from less than 100 kg N ha− 1 for the lower mineralization rate, to less than 300 kg N ha− 1 for the intermediate mineralization rate, to less than 500 kg N ha− 1 for the higher mineralization rates. The positive effect of the uptake rate on maize biomass is steeper for high mineralization rates.
Fallow periods allow for the constitution of important reserve of soil organic matter (and the N it contains) (Fig. 3). This reserve begins to decrease as soon as maize is sown due to mineralization, and maize grows absorbing the resulting mineral nitrogen. The soil is unable to fully regain its fertility between the two cropping cycles of the same year so that the yield of the second cropping cycle is lower than the first one. The biomass obtained from the first crop depressed from 185 kg N ha− 1 to 178 kg N ha− 1 for the second crop (Fig. 3). for the second crop. This is due to the quick mineralization of the soil organic matter stored during the fallow and the fact that the maize biomass (leaves and roots) added to the soil after the first maize cycle is not large enough to reach the quantity of soil organic matter reached at the end of the first fallow year.
Overall, maize biomass overall increased with the duration of fallows and decreased during successive years of maize cultivation (Fig. 4). There is an interaction between fallow duration and the number of successive years of cultivation. Hence, for one year of cultivation the yield remains high for successive fallow cycles and even increases for longer fallows. For 3 years of successive cultivation, the maize biomass decreases between fallow cycles for the shorter fallow, i.e. fallows no longer maintain fertility on the long term. For five years of successive cultivation, the yield decreases between fallow cycles whatever the length of the fallow.
Corn biomass varies greatly with the addition of N fertilizer and the effect of nitrification (Fig. 5). Without N fertilizer, the biomass of the two crops is almost identical. When we add fertilization (200 kg N ha− 1), we obtain for a crop under high nitrification a biomass of 242 kg N ha− 1 while the biomass under low nitrification is approximately 385 kg N ha− 1. We note that, maize biomass increases with the rate of N fertilizer application (Fig. 5). However, this increase is much stronger for a low nitrification rate during the cultivation than for a high nitrification rate. In other words, without mineral fertilization, low nitrification does not impact maize biomass, but the positive effect of low nitrification increases with the fertilization rate.
The legume-based fallow (improved fallow) has a much higher maize biomass than the other two fallow types, regardless of fallow duration (344, 450 and 657 kg N/ha respectively for fallow durations of 5, 10 and 20 years). The two other fallow types, resulted in almost similar maize biomass: low-nitrification fallow did not increase maize biomass sufficiently compared with the standard fallow (Fig. 6). For a 5-year fallow, we obtain a yield of 172 kg N/ha for low-nitrification and simple fallows. This yield increases slightly for a 10-year fallow (185 kg N/ha) and reaches 211 kg N/ha for a 20-year fallow
3.2 Discussion
Our results emphasized the particular influence of some of the mechanisms involved in N cycling, which overall confirms our hypotheses: (i) organic matter dynamics and mineralisation rates have a determining impact on soil fertility whatever the preference for NH4+ vs. NO3−; (ii) the inhibition of nitrification during the crop cycle increases the yield, but only in case of mineral N fertilization; (iii) long term fallow allows the building up a N reserve within soil organic matter, which maintains fertility as long as the fallow is long enough relatively to the number of successive cropping cycles; (iv) legume-based fallow increases soil fertility and crop yield much more than a nitrification-inhibiting one.
The initial results of the model confirmed the critical role of organic matter contributes in soil fertility replenishment and maintenance. According to Biaou et al. (2018), incorporating organic matter into the soil enhances its fertility through various mechanisms, one of which being considered in our model, i.e. the increase in the organic stock of mineral nutrients. Furthermore, organic matter mineralization rate plays a crucial role in enhancing maize yield (Thiebeau and Recous 2017). Current fertility requires a large enough stock of organic N and a sufficient instantaneous mineralization of this stock. Thus, in smallholder cropping systems with no mineral fertilization, soil fertility indeed depends on the past dynamics of organic matter accumulation during which inputs of organic N are higher than losses of N due to mineralization. This is partially due to mineral N being much more prone to losses than organic N.
Figure 2 clearly revealed that an increase in mineralization rate leads to a corresponding increase in maize yield, but only if a large soil organic matter stock has built up beforehand. The combined influence of absorption rate and NH4+ preference on maize growth biomass decreases with an increase in mineralization. In cases where the absorption rate is very high, the quantity of organic matter and mineralization rate become the decisive factors for successful cultivation of maize. Our results are in accordance with the empirical study of Cobo et al. (2002) that showed a strong correlation between organic matter decomposition rate, nutrient release, and the subsequent increase in yield due to nutrient absorption by crops. However, it is worth noting that mineralization, nutrient absorption and crop exportation necessarily gradually decrease fertility in a system without fertilization.
The preference for NH4+ vs. NO3− did not strongly impact maize biomass (Fig. 2) within a large range of preferences around 0.75 (i.e. 75% of N absorption as NH4+). Nevertheless, the yield strongly decreases for extreme values of preference and the lower the mineralization and the higher N absorption the narrower the range of preferences allowing for the maximal yield. These results are supported by previous modelling results (Boudsocq et al. 2009) showing that intermediate preferences for NO3− vs NH4+ with a preference for NH4+ increases primary production. Indeed, absorbing both NO3− and NH4+ prevents the crop to deprive itself of one of the two resources, and absorbing a little more NH4+ tends to reduce nitrification flows and subsequent losses by leaching and denitrification (Boudsocq et al. 2012; Konaré et al. 2019). While our study was based on constant preference values, it is important to acknowledge that the plant preference for a particular form of mineral N depends on many factors. Defining plant preference for NH4+ vs. NO3− can indeed be challenging, as it hinges on interactions between multitude of complex and dynamic environmental and physiological factors (Britto and Kronzucker 2013). These include soil chemistry, temperature, pH levels, plant genotype and stage of plant development (McFee and Stone 1968). Wang et al. (2021) demonstrated that maize prefers NH4+ in acidic soils, whereas Zhang et al. (2019) found that maize prefers NH4+ in neutral soils (subtropical humid monsoon climate). Nevertheless, further research is required in this area to better understand plant preference between NO3− and NH4+.
Like in past studies, our results confirmed that long fallow periods are required to maintain fertility in the absence of fertilization. For example, for 5 successive cropping cycles involving at least 20 years fallows are necessary for the N sustainability of the production. This is indeed due to the fact that exportations of N with the crop must be compensated, and that fallows allow for building up again fertility by stopping exportation and accumulating atmospheric inputs of N through dry and wet depositions that are absorbed by fallow vegetation and accumulate in the soil as organic N. Overall, our results confirm finding from empirical studies suggesting that the duration of fallow periods is important in restoring soil fertility (Atchada et al. 2019). In shifting cultivation systems, fallow duration significantly influences soil and vegetation dynamics, as observed in Fig. 4, which revealed that maize yield increases with the length of fallow periods (Toky and Ramakrishnan 1981; Swamy and Ramakrishnan 1988). However, the effectiveness of fallows decreases as cropping periods extend. To maintain the positive impact of fallows, it is necessary to increase the duration of the fallow period. This is confirmed by Kouelo (2015) who recommended a minimum fallow period of 30 years to restore degraded soils in sub-Saharan Africa. However this seems unrealistic as the need to satisfy food security of the growing population (Gafsi 2006; Abramova 2022) in sub-Saharan Africa makes it difficult to maintain such a long fallow period.
In our model, low nitrification during the cropping cycles increases maize yield only when mineral fertilization is used and this effect increases with the amount of N fertilizer (Fig. 5). In the absence of mineral fertilization, the possibility of NO3− leaching is low as maize absorbs a significant amount of NH4+ and NO3− so that the amount of available mineral N remains low all time. In this case, the nitrification rate (low or high nitrification) has minimal impact on maize biomass (Fig. 5). However, with inputs of mineral N fertilizer, the stock of mineral N increases transiently. In this case, a low nitrification rate reduces the quantity of NO3−, thereby minimizing losses through leaching and denitrification. These reduced N losses increase the availability of mineral N, in the form of NH4+, which promotes better plant growth and increases maize biomass. These results confirmed empirical results (Hu and Schraml 2014; Zhang et al. 2015a; Karwat et al. 2017) that showed that synthetic nitrification inhibitors with fertilizers reduce N losses and increase N availability, while maintaining good yield (Hu and Schraml 2014; Muller et al. 2023). However, our results seem to contradict findings by Boudsocq et al. (2012) and Konaré et al. (2019) suggesting that the inhibition of nitrification by savanna grasses strongly increases grass biomass production without any human fertilization. Two mechanisms likely explain the apparent discrepancy between these modelling results. First, our study focuses by definition on the transient dynamics of N cycling during a few cultivation cycles, whereas Boudsocq et al. (2012); Konaré et al. (2019) focused on long-term dynamics of ecosystems (equilibrium property of models). Such transient dynamics probably do not let enough time for the building up of N stocks through the repeated reduction of losses of NO3−. Second, the relatively high productivity of maize modelled requires a high rate of mineral N uptake by the crop. This reduced the availability of mineral nitrogen without fertilization, which may contribute to the lack of effect of low nitrification (Fig. 5).
According to Subbarao et al. (2012), sustainable agriculture requires the development of production systems incorporating new crop varieties capable of regulating nitrification and improved agronomic practices to reduce the leakage of N throughout the N cycle, a critical requirement to increase food production sustainably while mitigating adverse environmental effects. While inhibiting nitrification is viewed as very promising avenue to increase agriculture sustainability (Subbarao et al. 2012; Coskun et al. 2017), our model revealed some limits to these prospects. Rather logically, the availability of mineral NH4+ should be high enough and during a period sufficiently long for the nitrification inhibition to have an impact on yield. We also suggest that reduced nitrification would be more effective in agricultural systems with high mineral N inputs than in traditional subsistence agriculture without mineral fertilization.
Our results showed that the best way to improve the effect of fallows on N fertility is to grow N-fixing legumes in the fallow (Fig. 6). This confirms already well-established empirical results by Soro et al. (2015) that demonstrate the advantages of legume-based fallows even with short fallows (Williams et al. 2022). On the contrary, implementing a low-nitrification fallow does not significantly impact N fertility. This is likely explained, as for the low effect of a low nitrification rate during cropping cycles (see above), by the low availability of mineral N during the fallow (results not show). The impact of nitrification inhibition during fallow on yield and fertility remains largely unexplored experimentally. However, our results seem to be in contradiction with results showing that perennial savanna grasses, improve fertility probably through the inhibition of nitrification (Yé et al. 2017). A possible explanation is that we have used in our simulations a high value for mineral N absorption rate by fallow vegetation, while a lower absorption would increase mineral availability and increase the impact of reducing nitrification. Investing further this issue would require better describing N dynamics and the availability of NO3− and NH4+ during fallows.
Most agricultural system models are complex models that provide synthesis and quantification to assess the effects of water, soil, crops, trees, management practices and climate on the sustainability of agricultural production and to ensure food security (Dupraz et al. 2019; Burgess et al. 2019; Ahmed et al. 2022). On the contrary, our study was carried out using a fairly simple yet very general model, offering significant flexibility to consider key factors influencing N dynamics within the ecosystem. It is important to note that the model is not intended to lead to precise quantitative predictions, but rather to enhance our understanding of the system and to allow general predictions that are relevant to agriculture. To enhance the quantitative predictive accuracy of the model, we may consider incorporating additional compartments and mechanisms. For example, the absorption of mineral N, and the underlying complex plant-soil interactions, could be modelled using a function allowing for non-linearity between absorption, mineral stocks and plant biomass. Similarly, the preference for NO3− vs. NH4+ is probable not constant and influenced by many factors (Britto and Kronzucker 2013). These mechanisms could also be added in the model. Nevertheless, the generality of our model guarantees that our results are likely applicable to other crops and many fallow-based subsistence cropping systems from around the world. Up to our knowledge, it is also the first modelling effort analyzing the potential effects of biological nitrification inhibition on the sustainability of agriculture, in a context where N fertility and fertilization are at the heart of many sustainability issues (Zhang et al. 2015b).