Open access peer-reviewed chapter
Abstract
Camellia oleifera Abel. is one of the four woody edible oil trees around the world, which is also an important economic species in subtropical China. It is mainly cultivated in subtropical region, where the soil constrains the yield of C. oleifera oil due to its low fertility and pH. Thereby, intensive management including fertilization practice, especially intensive nitrogen (N) input, has been developed as a vital way to enhance oil yield in C. oleifera plantations. However, excessive nitrogen input increases soil nitrous oxide (N2O) emissions and soil acidification, limiting sustainable development of economic forests. As one of the important greenhouse gases, N2O is 265 times greater than carbon dioxide in global warming potential on 100-year scale. To mitigate soil N2O emissions and soil acidification, soil amelioration, including applications of biochar, nitrification inhibitors, and urease inhibitors, played an important role in sustainable management of C. oleifera plantations. This chapter reviewed soil nitrogen cycling, N2O emissions, and soil amelioration in C. oleifera plantations, which will benefit the sustainable management of C. oleifera plantations and hence the development of C. oleifera industries.
Keywords
- Camellia oleifera
- biochar
- nitrification inhibitor
- soil amelioration
- sustainable forest management
- urease inhibitor
1. Camellia oleifera
Specially,
As a typical economic tree, intensification such as water management, fertilization, and trimming takes an important part in the management of
2. Challenges
Fertilization is the major way of intensive management, efficiently improving the yield of oil in
2.1 Nitrous oxide emissions
Nitrous oxide, as the major ozone-depleting substance [10], has been recognized to be an important greenhouse gas. Especially, the potential of N2O for global warming is 265 times than that of carbon dioxide [11]. The concentration of N2O is ranging from 270 ppb in pre-industrial period to 329.9 ppb in 2017 [12].
Soil systems contributed the largest source of N2O emissions (13 Tg N2O-N yr−1), of which human activities accounted for 54% [13]. Nitrogen input such as N deposition and N fertilization often increased N2O emissions and altered the process of N transformation [14, 15, 16, 17]. Generally, soil N2O emissions had a positive and linear relationship with N input [18]. About 120 Tg N was contributed by human activities per year [13]. Therefore, intensive N input often leads to high emissions of soil N2O [19].
2.1.1 Nitrification and denitrification
Nitrification and denitrification are the two main pathways of N2O emissions (Figure 1) [20, 21, 22], which produced global 70% soil N2O emissions [13].
Figure 1.
Nitrification- and denitrification-related pathways [
2.1.2 Influence factors
Soil N2O emissions can be influenced by soil environmental factors such as soil moisture, temperature, oxygen (O2), and pH condition [23, 24].
2.1.2.1 Soil moisture
Soil moisture is a vital factor that affects soil N2O emissions. Generally, soil N2O emission rates reached the peak when soil water-filled pore space (WFPS) was 60–70% [25]. For example, soil N2O emissions were significantly higher under 60% WFPS conditions than that under flooded conditions [26].
2.1.2.2 Soil temperature
Effects of soil temperature on N2O emissions were more complex than that of soil moisture. For example, warming increased soil N2O emissions from boreal peatland [27] and alpine meadow [28]. Soil N2O emissions had an exponential increased relationship with incubation temperatures [29]. A significant positive correlation was presented in N2O emissions and soil temperature from different soil types (paddy, orchard, forest, and mountain) [30]. Although warming did not affect soil N2O emissions from northern peatlands, it suppressed N2O emissions under N addition conditions [31]. By contrast, the effects of warming on soil N2O emissions from alpine meadow soil were not observed [32]. Consistently, no significant increase of soil N2O emissions was found with increasing incubation temperatures [33]. Previous study reported that soil moisture and temperature can explain 86% of soil N2O emissions [34].
2.1.2.3 Soil O2 concentration
Soil O2 concentration was closely related with soil moisture and soil mechanical composition. Generally, soil with higher water content and larger clay fraction had lower soil O2 concentrations. Lower soil O2 concentrations mainly promoted soil N2O emissions via denitrification [20, 35]. The production of N2O and NO was increased when O2 concentration decreased from 21% to 0.5% in a laboratory study [36]. Similarly, field study reported that soil N2O emissions increased with increasing soil O2 concentrations in wetland [37].
2.1.2.4 Soil pH
pH played an important role in the activity of microbes [38]. Indeed, soil acidification [39] and soil pH amelioration [40] significantly influenced soil N2O emissions. However, other researchers reported that there was no significant correlation between N2O emissions and pH [41].
pH influenced the activity of nitrification- and denitrification-related enzymes [42]. Generally, soil acidification increased N2O emissions [42]. Compared with ammonia-oxidizing bacteria (AOB), ammonia-oxidizing archaea (AOA) were higher in activity and resistance from acid soil [43]. However, the domination of AOB was increased by increasing soil pH [44]. Additionally, archaeal
2.1.3 Nitrous oxide emissions from Camellia oleifera plantation soils
Our previous field study (1 year) found that soil N2O emissions were 92.14 ± 47.01 mg m−2 in control treatment and were 375.10 ± 60.30 mg m−2 in fertilization treatment (400 kg NH4NO3-N ha−1) from
2.2 Soil acidification
Acid soil (pH < 5.5) as a main soil type covers about 30% free ice land [46]. However, soil acidification has been becoming more and more serious [47]. Soil acidification should be taken into consideration due to its constraint in the sustainable development of agricultural sector [48]. In China, soil pH (except alkaline soils at pH 7.10–8.80) from crop fields reduced by 0.13–0.76 during the year 1980–2000 [49]. For example, soil pH (surface layer) decreased by 0.30 units from 1981 to 2012 in Sichuan Province, China [47].
With a long cultivation history,
Soil acidification from
2.2.1 Precipitation
Long-term precipitation increased the loss of base cations such as Ca2+, Mg2+, K+, and Na+, reducing the soil pH buffering capacity. In addition, long-term precipitation promoted the accumulation of Al3+ and Fe3+ in soil, which could further hydrolyze to Fe(OH)3 or Al(OH)3 and release 3H+.
2.2.2 Plant physiology
When plant roots absorb a NH4+ from soil, an H+ will release into soil; in turn, absorbing a NO3− from soil will release an OH− into soil [53].
Organic acid (R▬COOH) from root exudates can release an H+ after hydrolysis. In addition, anions of organic acids (e.g., citric acid and malic acid) can chelate with Al3+ in the soil and inhibit the root system that absorbs Al3+, alleviating Al3+ toxicity to plant growth [48, 54, 55].
Plants such as
2.2.3 Microbial-mediated nitrification
For example, NH4+ transfers to NO3− along with the 2H+ release (NH4+ + 2O2 → NO3− + H2O + 2H+) [53]. AOB, AOA, and fungi can participate in the process of nitrification [20]. Nitrification includes the pathway of ammonia oxidation to hydroxylamine, the pathway of hydroxylamine oxidation to nitrite, and the pathway of nitrite oxidation to nitrate (Figure 1) [58]. Ammonia can be oxidized by AOA or AOB to hydroxylamine via ammonia monooxygenase (
2.2.4 Oxidation of sulfur-containing organics
Oxidation of sulfur mineral, for example, oxidation of FeS2, will produce 2H+ (2FeS2 + 7O2 + 2H2O → 2Fe2+ + 4SO42− + 4H+).
Oxidation of sulfur-containing organics will release 4H+ (2Organic-S + 3O2 + 2H2O → 2SO42− + 4H+).
2.2.5 Intensive nitrogen fertilization
Intensive NH4+ input can replace the base cations such as Ca2+, Mg2+, K+, and Na+ and accelerate leaching, reducing the pH buffering capacity of top soil [59]. Hydrolysis of soil NH4+ will generate NH3 (gas) and consume an OH− (NH4+ + OH− = NH3↑ + H2O) [60].
Acidic fertilizers such as Ca(H2PO4)2 will gradually release H+, hence increasing soil acidification (Ca(H2PO4)2 → CaHPO4 + H3PO4, H3PO4 → H+ + H2PO4 → 2H+ + HPO42− → 3H+ + PO43−).
2.2.6 Acid deposition
Acid deposition (water-soluble acid gases such as CO2 and sulfur dioxide) and N deposition (especially NH4+-N) increased soil acidification [51]. Precipitation with H+ can replace the soil base cations such as Ca2+, Mg2+, K+, and Na+, which directly reduce the soil pH buffering capacity [51].
2.2.7 Other factors
For example, deforestation and other land uses can reduce litter accumulation in surface soil, hence declining the accumulation of base cations such as Ca2+, Mg2+, K+, and Na+ that generate from litter decomposition [61].
2.3 Effects of soil acidification on nitrous oxide emissions
Acid soils have been facing an increased risk of acidification due to human activities, especially intensive N fertilization [47, 49, 62]. For example, after 6 years of application of 600 kg Urea-N ha−1 yr−1, soil pH was significantly decreased (soil pH in control and fertilization treatment was 5.1 and 4.9, respectively) from a tea plantation in Yixing City, Jiangsu Province, China [63]. A meta-analysis of 1104 field data showed that a negative correlation between soil N2O emissions and pH (3.34–8.7) (N2O-N = −0.67x + 6.55,
The mechanism of soil acidification on the stimulation of soil N2O emissions is complex, which may include (but not limited to) the following points.
2.3.1 Chemical decomposition of nitrous acid
Under acidic conditions, pH < 5.5, NO2− (HNO2, pKa = 3.3) will naturally decompose into NO and/or NO2 (3HNO2 ⇌ 2NO + HNO3 + H2O or 2HNO2 ⇌ NO + NO2 + H2O) [65]. Soil NO can be further transformed to N2O with Fe2+ when it was not escaping soil [65].
2.3.2 Shifts in microbial communities and abundance
Generally, the abundance of AOB was lower in soil pH < 5.5 than that in neutral soil pH. Here, nitrification was weak and almost disappears at soil pH < 4 [66]. However, AOA could mediate the process of ammonia oxidation in extremely strong acidity soil (pH: 4.2–4.47) [43]. Another study reported that the abundance of AOB was positively correlated with pH (
In acid soils, the activity of N2O reductase was inhibited, leading to higher N2O emissions in lower soil pH [72]. Indeed, there was a positive correlation between the abundance of
2.3.3 Microbes increased resistance to soil acidification
Laboratory study showed that the potential of soil N2O emissions was increased with decreasing pH (soil pH ranging from 2.96 to 6.26) from tea plantations in Japanese [78]. In addition, higher soil N2O emissions and lower abundance of
3. Sustainable forest management
Soil amelioration (e.g., application of lime, biochar nitrification inhibitors, and urease inhibitors) plays an important role in mitigation of soil acidification and N2O emissions.
3.1 Lime
Lime as an ameliorant was often used to amend acid soils in southern China due to increasing soil pH. It can relieve the toxic effect of soil Al3+ on plant growth by reducing soil exchangeable H+ [81]. Lime addition increased soil pH and salt saturation [82]. In addition, application of lime can reduce soil N2O emissions [40]. For example, under 60% WFPS or flooded conditions, dolomite addition at medium- or high-dose levels (1 or 2 g kg−1 soil) can reduce N2O emissions and increase the transcription of
3.2 Biochar
Biochar was stable in the soil from Amazon basin of Brazil, and biochar input improved soil fertility [85]. This discovery accelerated the development of technologies for biochar application in soil amelioration.
Biochar is a carbon (C)-rich solid material by pyrolyzing of organic biomass such as crop straw, forestry by-products, urban waste, industrial by-products, animal manure, and urban sludge at low oxygen and high temperature (250–700°C) condition [86]. Biochar has been characterized by a high pH, specific surface area, degree of aromatization, and porosity. In addition, biochar is rich in C-containing functional groups (e.g., C–H, C–O, C=C and C=O) and relatively stable organic C. The physicochemical properties of biochar were mainly determined by pyrolysis temperature [87].
Presently, biochar was widely used as a soil ameliorant in agriculture and forestry field. For example, our previous studies reported that
Carbon recalcitrance of biochar can increase soil C pool. The potential of biochar in mitigation of greenhouse gas emissions was 1.0–1.8 Pg CO2-Ceq yr−1 [89].
Biochar had excellent physicochemical characteristics in soil nutrient retention and utilization [90, 91] and water conservation [92]. Additionally, biochar can increase the plant resistance to Al3+ toxicity [81], the clone of arbuscular mycorrhizal fungi, and crop yield [93, 94]. It can decrease continuous cropping obstacles such as root-knot nematode [95] and
Ralstonia solanacearum [96].Biochar is rich in macro- and microelements [97], which can reduce the dosage of fertilizer.
3.2.1 Effects of biochar on soil nitrous oxide emissions
The physicochemical properties of biochar and soil can interactively influence soil N2O emissions [98]. However, the effects of biochar on soil N2O emissions varied, including positive effects [99], negative effects [100], and no effects [101].
Biochar addition increased soil N2O emissions with the release of N from biochar [102]. By contrast, biochar reduced soil N2O emissions with (1) increased NO3−-N immobilization [103]; (2) increased copy numbers of
3.2.2 Effects of biochar on soil pH buffer capacity
Biochar that increased soil pH buffer capacity may predominantly correlate with biochar riches in oxygen-containing functional groups in surface. The anions of weakly acidic functional groups can associate with H+, hence increasing soil pH. Meanwhile, exchangeable base cations can release into the solution, thus increasing soil pH buffer capacity [107, 108]. In addition, soluble silicon (Si) such as H3SiO4− (present at a high pH) can combine with H+ and generate H2SiO3 precipitation [107, 108].
3.3 Nitrification inhibitor
Nitrification inhibitors are a class of organic compounds that can inhibit the activity of nitrifying bacteria.
Nitrification inhibitors, especially synthetic nitrification inhibitors (e.g., dicyandiamide (DCD) and 3,4-dimethylpyrazole phosphate (DMPP)), were widely used in agriculture for improving N use efficiency. Ammonia-oxidizing bacteria and AOA are the major microbial communities in nitrification and denitrification, and both contain
Ammonium N can be adsorbed by soil colloids, while soil NO3−-N (the end product of nitrification) easily can be leached to groundwater by precipitation. In addition, microbial-mediated nitrification is closely related with soil N2O emissions [20, 21, 22]. Nitrification inhibitors can effectively inhibit soil nitrification, slowing the transformation of NH4+-N to NO3−-N and hence reducing the NO3−-N leaching and N2O emissions.
An evaluation from 62 field studies showed that although nitrification inhibitors increased 20% NH3 emissions, they reduced 48% inorganic N leaching, 44% N2O emissions, and 24% NO emissions and increased 58% plant N utilization, 9% grain yield, 5% straw yield, and 5% vegetable yield [115]. Consistently, other studies evaluated that nitrification inhibitors decreased by 38% [116], 50% [117], or 73% [118] N2O emissions and decreased by 0.3 t CO2e ha−1 yr−1 [119]. Similarly, DCD did not increase crop yields but reduced 35% N2O emissions [120]. A meta-analysis showed that DCD rather than DMPP significantly increased 6.5% crop yield as well as DCD and DMPP decreased N2O emissions by 44.7% and 47.6%, respectively [121].
Therefore, application of nitrification inhibitors could reduce N2O emissions and mitigate environmental pollution after intensive N inputs.
3.4 Urease inhibitors
Urease inhibitors are a class of compounds that can slow soil urease activity (Figure 2). Addition of urease inhibitors after urea input can inhibit the hydrolysis of urea via inhibiting the activity of urease, hence reducing NH3 volatilizations and N2O emissions. Additionally, the application of urease inhibitors also contributes to increase N utilization efficiency and reduce NO3−-N leaching.
Figure 2.
The chemical equation of urea hydrolysis with urease catalysis.
Urease, a Ni-copper enzyme, has two Ni−O bidentate ligands, specifically catalyzing urea into NH3 and CO2. Urea only can bind with one specific Ni−O ligand of urease, but NBPT can bind with two Ni−O bidentate ligands of urease and generate a tridentate ligand [122], hence inhibiting the activity of urease.
Presently, a meta-analysis reported that a nonlinear response was presented in soil NH3 volatilizations and N input [123]. Application of NBPT can effectively inhibit NH3 volatilizations. For example, 530 mg NBPT kg−1 urea treatment delayed NH3 volatilizations and decreased accumulation of NH3 volatilizations compared with the control treatment. NH3 volatilizations were linearly related with the NBPT dosage in the range of 0–1000 mg NBPT kg−1 Urea (0, 530, 850, 1500, and 2000 mg NBPT kg−1 Urea) [124]. Other study reported that NBPT increased 27% oat yield and 33% crop N uptake [120].
The effects of NBPT on N2O emissions were controversial. For example, NBPT can reduce 80% N2O emissions [117]. No effects of NBPT (0.07%, NBPT/Urea-N, w/w) on N2O emissions were observed [125]. Similarly, there was no change of N2O emissions with NBPT (250 mg NBPT kg−1 Urea) addition from urea-fertilized (50 kg Urea-N ha−1) soil [126].
Additionally, NBPT can reduce N2O emissions from alkaline soils but has no effects on acidic soils [127], which indicated that pH plays a key role in the regulation of NBPT effects on N2O emissions. Further laboratory study showed that NBPT inhibited nitrification, stimulating N2O emissions from alkaline soils (pH 8.05) but not affecting N2O emissions from acid soils (pH 4.85). This finding suggested that the effect of NBPT on soil N2O emissions is not only influenced by pH but also by other unknown factors [127].
Generally, urease inhibitors correlated with nitrification inhibitor could mitigate N2O emissions. A meta-analysis showed that urease inhibitors and nitrification inhibitors interactively reduced 30% N2O emissions [116]. For example, a field study reported that the combination of NBPT (0.3%, NBPT/Urea-N, w/w) and DCD (0.3%, DCD/Urea-N, w/w) reduced 32.1% soil N2O emissions with the addition of 519 kg Urea-N ha−1 from banana plantation, but did not affect the yield of banana [128].
4. Sustainable management in Camellia oleifera plantations
Our previous incubation study found that although biochar application increased N2O emissions, DCD addition decreased soil N2O emissions under urea fertilization from
Figure 3.
The seed yield of
5. Conclusions
Soil acidification, especially induced by N fertilization, will inhibit the activity of N2O reductase and increase the abundance of N2O-producing fungi as well as the acid resistance of N2O-producing microorganisms, hence the ratio of N2O/(N2O + N2). In addition, NO2− will generate NO under soil pH < 5.5 condition, which will further transform into N2O. Under the background of global acidification, the soil from
Acknowledgments
The National Natural Science Foundation of China (grant number: 41967017 and 41501317), Jiangxi and China Postdoctoral Science Foundation (grant number: 2017M106153 and 2017KY18), and Jiangxi Education Department (Project Number: GJJ160348) support this work.
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