Climate warming and nitrogen deposition had no significant interaction effect on litter decomposition of subtropical mountain forest in Southwest China

doi:10.21203/rs.3.rs-3211295/v1

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Research Article

Climate warming and nitrogen deposition had no significant interaction effect on litter decomposition of subtropical mountain forest in Southwest China

https://doi.org/10.21203/rs.3.rs-3211295/v1

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published 11 Apr, 2024

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Background and Aims

Litter decomposition is essential in the global carbon cycle, which is affected by environmental factors. Under the context of global warming and increasing nitrogen deposition, the responses of litter decomposition are unclear in subtropical forests.

Methods

Here, we carried out an interaction experiment of warming (3.3 ^oC) and nitrogen addition (50 kg ha^− 1 yr^− 1) and a recovery experiment (remove warming and nitrogen addition treatments) in a subtropical evergreen broad-leaved forest for four years in total.

Results

The results showed that during the treatment period, warming, nitrogen addition, and their combination significantly decreased litter decomposition according to the k values by 35.8, 23.0, and 42.2%, respectively. However, there was no significant interaction between warming and nitrogen addition. Litter decomposition rates quickly returned to the same level during the two-year recovery period. Variance partitioning analysis (VPA) and covariance analysis (ANCOVA) simultaneously confirmed that warming and nitrogen addition significantly affected litter decomposition. VPA and ANCOVA also showed that soil temperature and water content significantly contributed to the decomposition rate in the treatment period.

Conclusions

This study suggests that warming and nitrogen addition significantly inhibit litter decomposition, but this effect is not sustainable.

Litter decomposition

Climate warming

Nitrogen addition

Soil environmental factors

Initial decomposition degree

Litter decomposition is essential in the global carbon cycle (Daan et al. 2018). Current studies suggest that litter decomposition is mainly driven by microorganisms, which are also influenced by climate, litter quality, soil environmental factors, and the initial priming effect (Ge et al. 2013; Liu et al. 2023). At present, the continuous rise in temperature and atmospheric nitrogen deposition caused by human activities has become the main content of climate change (Liu et al. 2011; De Frenne et al. 2021), which significantly affects the process and function of ecosystems (Grimm et al. 2013). Therefore, increasing studies have focused on the effects of warming and nitrogen addition on litter decomposition (Henry and Moise. 2015; Ma et al. 2011). However, the relevant research has no clear conclusion.

Previous studies have shown that the effect of warming on the litter decomposition rate may be easily affected by water conditions in the environment (Butenschoen et al. 2011; Gorecki et al. 2021). A certain degree of warming can increase the activity of microorganisms, thereby promoting litter decomposition (Aerts. 2006). However, the decrease in soil water content caused by warming will inhibit microbial activity, which limits litter decomposition (Hanna et al. 2014). Previous studies have confirmed that in some ecosystems with adequate water conditions, such as rivers, wetlands, and peatlands, warming usually promotes litter decomposition (Ferreira et al. 2013; Gao et al. 2022; Migliorini. 2020). In contrast, in some arid ecosystems, warming tends to inhibit litter decomposition (e.g., grasslands and plateaus) (Chuckran et al. 2020; Penner and Frank, 2018). Other studies have shown that warming has greater variability in litter decomposition in mountain forest ecosystems with complex structures (Pietsch et al. 2019). Some studies have reported that warming will promote litter decomposition in forest ecosystems (Liu et al. 2015; Liu et al. 2017), while others have said that warming will significantly inhibit litter decomposition (Li et al. 2022; Liu et al. 2022). Therefore, warming effects may be related to the complex hydrological processes of mountain forest ecosystems (Jones et al. 2022).

Similar to the effect of warming on decomposition, the response of external nitrogen addition to litter decomposition is also highly uncertain (Zhang et al. 2018). Previous studies have shown that nitrogen addition may promote litter decomposition (Song et al. 2018; Zhang et al. 2019), inhibit litter decomposition (Peng et al. 2022; Zhang et al. 2022), or have no effect (Růžek et al. 2021). It is speculated that the different effects of nitrogen addition on litter decomposition may be closely related to the local atmospheric nitrogen deposition level, litter quality, and nitrogen addition concentration (Knorr et al. 2005; Schulte-Uebbing and de Vries. 2018). In addition, nitrogen addition may also lead to changes in soil water content by influencing the water absorption efficiency of dominant plant roots, thus indirectly influencing litter decomposition (Henriksson et al. 2021). Therefore, the effects of warming and nitrogen addition on the litter decomposition rate may be affected by various factors and have great uncertainty (Zou et al. 2022). Current research on forest ecosystems focuses on the impact of individual factors on litter decomposition (Xia et al. 2013). The interaction between warming and nitrogen addition in litter decomposition is rarely involved (Henry and Moise. 2015; Moise and Henry. 2014).

Subtropical forests play an essential role in the global terrestrial ecosystem carbon cycle, with a wide range and a large amount of litter (Tan et al. 2011; Wu et al.2014). Currently, subtropical forests in southwestern China suffer from climate warming and high nitrogen deposition (Chen and Mulder. 2007; Zhou et al. 2022). However, few studies have explored the response of litter decomposition to warming and nitrogen addition in similar regions (Zhang et al. 2017). The subtropical evergreen broad-leaved forest in Ailao Mountain of China has an altitude of more than 2400 meters, an average annual temperature of 11.3 ^oC, and abundant rainfall (Wu et al. 2014). Based on the above, we hypothesized that 1) warming and nitrogen addition promote litter decomposition, and there is a significant positive interaction; and 2) warming and nitrogen addition have sustained effects on litter decomposition. We carried out an experiment of warming and nitrogen addition to test the hypotheses in a subtropical forest. We also conducted a leaf litter decomposition experiment after the treatments were terminated (recovery experiment) to test the sustainability of the treatments.

2.1 Study site description

This study was conducted at the Ailaoshan Station for Subtropical Forest Ecosystem Studies (24°32'N, 101°01'E; 2480 m above sea level), which is located in Jingdong County, Yunnan Province. It is an old-growth subtropical evergreen broad-leaved forest that has been deeply studied (Tang et al. 2007). It is characterized by a subtropical mountain climate with an annual average temperature of 11.3 ^oC and annual precipitation of 1840 mm (Wu et al., 2014). There is an obvious rainy season from May to October every year. The forest soil is rich in nitrogen and acid (pH < 5) (Chan et al. 2006). The main tree species of the forest are Machilus gamblei, Castanopsis wattii, and Hartia sinensis and the litterfall is 864 g m^− 2 year^− 1 (Wu et al. 2014).

2.2 Experimental design and setup

In this study, we selected 12 subplots (1×1 m) in a 30×40 m experimental plot. We conducted two factors with two levels of experiments: control (CK), warming (W), nitrogen addition (N), and warming together with nitrogen addition (WN). Each treatment had three repeats and contained treatment and recovery periods. Three lamps with a length of 90 cm were set up in the warming area for warming treatments. The lampshade was 100 cm and 1.2 m from the ground. The three lights form a triangle tilted and fixed at a 30^o angle to the bottom to collect heat. We added NH₄NO₃ solution at 50 kg N ha^− 1 yr^− 1 for nitrogen treatments, divided into 12 months per year. Therefore, we weighed 1.19 g NH₄NO₃, dissolved it in 500 ml ultrapure water, and sprayed it evenly into the subplots once a month. We also spread the same volume of ultrapure water for the subplots with no nitrogen addition.

We collected fresh litter leaves from the subtropical forest during the dry season from October to December 2017, air-dried them, and dried them in an oven at 60 ^oC for 72 hours to constant weight. We prepared 12 litter bags (2 mm mesh, size 10 × 15 cm) for each subplot with an initial mass (M_i) of approximately 4 g. After removing the visible litter, we placed the litter bags on the soil surface 10 cm apart on December 31, 2017. Here we intend to carry out the decomposition experiment for four years. Therefore, the decomposed litter samples were collected three times per year. Litter decomposition during the treatment period was terminated for many reasons, and a recovery experiment was conducted instead. We prepared 6 litter bags (2 mm mesh, size 15 × 30 cm) for each subplot with approximately 14 g initial mass for the recovery period (fresh litter leaves were collected from early January to mid-March 2021). The recovery experiment started on March 31, 2021, and lasted for two years.

2.3 Data collection and analysis

As mentioned above, we collected litter bags every four months for two years for a total of 6 times in the treatment and recovery period. After each collection of litter bags, we cleaned, washed, and dried them at 60 ^oC for 72 hours to constant weight to obtain the litter dry weight (M_f).

We calculated the decomposition coefficient (k value) during the study period using the following decay model (Olson, 1963).

$$RR=\frac{{M}_{f}}{{M}_{i}} \left(1\right)$$

$$RR={e}^{-kt} \left(2\right)$$

where RR is the residual rate, and t (yr) is the corresponding decomposition time.

We divided litter decomposition into six stages and calculated the stage decomposition rate (SDR, %/month).

$${SDR}_{s}=\frac{{RR}_{s-1}-{RR}_{s}}{{RR}_{s-1}} ÷{T}_{stage}÷12\times 100\text{\%} \left(3\right)$$

where is the stage number from 1 to 6, and T_stage is each stage time (yr).

Every month, we measured the soil temperature (^oC) and soil water content at a 5-cm depth using a digital thermometer (6310; Spectrum, Illinois, USA) and time domain reflectometry (MP-KIT; Beijing Channel, Beijing, China) three times from January 2017 to March 2023. We divided them into three periods: before treatment (2017.01-2017.12, background of each subplot), treatment period (2018.01-2019.12, treatment effect), and recovery period (2021.04-2023.03). The averages of each period were used to test the treatment effect, taking the before-treatment period as the background value and calculating the ratio value of the treatment and recovery period. We also divided the soil temperature and water content into six stages (SST and SSW) corresponding to the SDR in the treatment and recovery periods.

One-way ANOVA and Duncan’s test were used to test the difference in each period’s soil temperature and water content, ratio values, and k values among treatments. Two-way ANCOVA was used to test each period’s k values and SDR difference. Warming and nitrogen addition treatments were the fixed factors, and soil water content was the covariate for k values. SST, SSW, and initial decomposed degree (IDD) were the covariates for SDR. All the data passed the normality and homoscedasticity test. The effects of warming, nitrogen addition, IDD, SST, and SSW on the SDR were analyzed by variance partitioning analysis (VPA). The variance inflation factor (VIF) test was conducted, and all VIF values were less than 4. All statistical charts were generated using R software (version 4.3.0) and R Studio with the readxl, ggplot2, lubricate, ggpubr, car, gtsummary, agricolae, and ggbeeswarm packages.

3.1 Effects on soil microenvironmental factors

Warming and nitrogen addition did not change the seasonal variation in soil temperature and water content in this study (Fig. 1). There was no significant difference in temperature between all plots before and after treatment (Fig. 2a). Soil temperature had no significant difference among each plot before treatment, only in the treatment period, was the temperature of the warming group and the warming and nitrogen addition group significantly higher than that of the non-warming group, with a temperature increase of 3.3 ^oC (Fig. 2a, 2b). In contrast to different from the trend in temperature, the soil water content had high heterogeneity before treatment (Fig. 2c). Considering the heterogeneity of the initial water content, the soil water content in the treatment stage and recovery stage of the sample plot was compared with soil water content before treatment to show its change (Fig. 2d). The results showed that during treatment period, the soil water content in the W and WN plots was significantly lower than that in the CK and N addition plots (Fig. 2d). In general, soil temperature returned to the same level in all plots after removal of warming and nitrogen addition treatments (recovery period). Meanwhile, the soil water content of the other plots also recovered to similar levels except for the warming treatment plots.

3.2 Effects of warming and nitrogen addition on litter decomposition

In the treatment period, the differences in the litter residual rate increased with decomposition time (Fig. 3a). One-way ANOVA showed that all treatments (N, W, WN) significantly affected the litter decomposition rate in terms of k value and SDR in the treatment period, and the k values of CK, N, W, and WN were 0.422 ± 0.008, 0.325 ± 0.046, 0.271 ± 0.032 and 0.244 ± 0.039 (mean ± SD), and the mean values of SDR were 3.20 ± 0.12, 2.33 ± 0.28, 2.16 ± 0.45, and 1.85 ± 0.30%/month (mean ± SD), respectively. Where N, W, and WN treatments reduced the k values by 23.0, 35.8, and 42.2%, and reduced the SDR by 27.2, 32.5, and 42.2%, respectively (Fig. 3b, d). In the recovery period, there was no significant difference in K values and SDR between treatments (Fig. 3b, d). Two-way ANCOVA showed that warming and nitrogen addition had significant effects on the k values and mean SDR without considerable interaction, and the covariate had no significant effect (Table. 1). Further analysis also showed no difference after the warming and nitrogen addition treatments were terminated (Table. 1), suggesting the effects of warming and nitrogen addition on litter decomposition were not persistent.

Table 1

Covariance analysis of nitrogen, warming, their interactions and soil water content on litter decomposition rate (K value) and litter stage decomposition rate (SDR) during treatment and recovery periods.
Effect	Treatment period					Recovery period
Effect	Df	Sum sq	Mean Sq	F value	Pr	Df	Sum sq	Mean Sq	F value	Pr
Factor(N)	1	6.18	6.18	5.62	0.02	1	0.25	0.25	0.21	0.65
Factor (Warming) Factor(N):factor(Warming) Ts Ws IDD	1 1 1 1 1	10.47 1.79 38.47 17.35 2.90	10.47 1.79 38.47 17.35 2.90	9.51 1.63 34.93 15.77 2.64	0.01 0.21 < 0.001 < 0.001 0.11	1 1 1 1 1	0.61 1.45 67.81 30.38 33.65	0.61 1.45 67.81 30.38 33.65	0.51 1.22 56.91 25.50 28.24	0.48 0.27 < 0.001 < 0.001 < 0.001
Residuals	65	71.58	1.10			7	77.46	1.19

3.3 Factors affecting litter decomposition

To explore the factors affecting litter decomposition, we used the litter mean SDR, IDD, and the corresponding soil environmental factors for further analysis (Figs. 3c, 4). Two-way ANCOVA showed that the SDR significantly differed due to warming and nitrogen addition, and was affected by SST and SSW (Table. 2). Nevertheless, only SST, SSW, and IDD in the recovery period greatly influenced the SDR (Table. 2), suggesting that the effect of the treatments disappeared.

VPA showed that SST, warming, and SSW significantly contributed to the SDR by 36.7, 25.3, and 24.5% in the treatment period, respectively. IDD and nitrogen addition contributed 8.3 and 5.2%, respectively (Fig. 5a). In the recovery period, IDD significantly contributed to 60.4% of the SDR; SST and SSW contributed 20.9 and 7.1, respectively, without significance (Fig. 5b).

Table 2

Covariance analysis of nitrogen, warming, their interactions, Ts, Ws and IDD on litter stage decomposition rate (SDR, %) during treatment and recovery periods.
Factor	K values				SDR
	Treatment period		Recovery period		Treatment period		Recovery period
	F	P	F	P	F	P	F	P
N	8.9475	0.0202	0.1198	0.7394	10.5322	0.0141	0.2468	0.6345
Warming	31.3716	0.0008	1.0645	0.3365	17.8359	0.0039	0.6014	0.4634
N: Warming	2.9987	0.1269	0.4867	0.5079	2.6315	0.1488	0.8996	0.3745
Ws	0.0621	0.8103	1.2334	0.3034	0.5012	0.5018	0.6553	0.4449

Temperature is a critical factor affecting litter decomposition (Liu et al. 2023; Wu et al. 2022). In general, modest warming will to some extent promote the activity of microorganisms, thereby promoting litter decomposition (Liu et al. 2015). Previous research reported that an increase in soil temperature by 1.84 ^oC significantly promoted litter decomposition by 10.3% (Gao et al. 2022). Other studies also found similar results (Ferreira et al. 2013; Migliorini et al. 2020). However, the results of this study showed that the litter decomposition rate decreased significantly when the temperature increased by 3.3 ^oC (Fig. 2c, d), similar to previous studies (Li et al. 2022; Penner and Frank. 2018; Liu et al. 2022). We speculate that this may be related to the decrease of soil water content caused by warming. In this study, the soil water content under warming treatment decreased by 18.2%, which was consistent with earlier studies in forest ecosystems (Liu et al. 2015; Liu et al. 2017; Li et al. 2022). The decreasing soil water content may significantly inhibit the activity of microorganisms, thus inhibiting decomposition (Butenschoen et al. 2011; Liu et al. 2022b). Meanwhile, the dry heat stress caused by warming treatment may also indirectly inhibit litter decomposition by inhibiting the quantity and activity of soil animals (Barajas-Guzmán and Alvarez-Sánchez. 2003; Collison et al. 2013). VPA showed that soil temperature and water content positively were correlated with the litter decomposition rate. In contrast, the warming treatment was significantly negatively correlated with the litter decomposition rate (Fig. 5a). This further indicates that the inhibitory effect of the decrease in soil water content on litter decomposition may be much greater than the promoting effect of soil warming. Therefore, the impact of warming on litter decomposition in forest ecosystems depends on the combined effects of warming and the adverse effects of the changes in soil water conditions caused by warming (Ferreira and Canhoto. 2013; Canessa et al. 2020;).

Previous studies suggested that nitrogen addition could improve litter quality (with a lower C/N ratio), thereby promoting litter decomposition to a certain extent (Jiang et al. 2014; Tian et al. 2018). However, similar conclusions were not reached in our study. We speculate that this may be closely related to the concentration of nitrogen addition (Tu et al. 2014). Previous studies have shown that a low concentration of nitrogen addition promotes litter decomposition, while a high concentration inhibits litter decomposition (Knorr et al. 2005). In this study, the nitrogen concentration was 50 kg ha^− 1 yr^− 1, which is approximately 3 to 4 times the local atmospheric nitrogen deposition (Liu et al. 2002). Previous studies have shown that a continuous high concentration of nitrogen addition might alter microbial community composition and function (Wu et al. 2019), inhibit the activity of catabolic enzymes (Jiang et al. 2014), and reduce the amount of nitrogen they need to take up from litter, thereby inhibiting litter decomposition. In addition, the addition of high nitrogen concentrations may also decrease soil arthropod group number, individual number, and diversity, thereby inhibiting litter decomposition (Tie et al. 2021).

In this study, warming and nitrogen addition significantly inhibited litter decomposition (Fig. 3b, d). However, there was no significant interaction between warming and nitrogen addition (Table. 1). which differs from previous research conclusions in grassland ecosystems and wetlands (Ma et al. 2011; Tao et al. 2019). It is speculated that this may be related to the decreased soil water content caused by the warming treatment (Chuckran et al. 2020). Under low soil water content, the litter decomposition rate is significantly inhibited. The inhibitory effect of nitrogen addition was weakened (Liu et al. 2022). Therefore, the inhibition effect of WN on litter decomposition does not show a noticeable amplification effect compared to warming and nitrogen addition alone. In addition, it was observed that the soil water content of the initial WN plot was also relatively lower than that of the other treatment groups (Fig. 2c), which may further weaken the interaction. Therefore, soil water content variation may influence the interaction between warming and nitrogen addition on litter decomposition.

The experimental results in the recovery period of this study showed that the decomposition rate of litter rapidly recovered to the level of the control. Meanwhile, soil environmental factors recovered well after removing the warming and nitrogen addition treatments (Fig. 3a, c), suggesting no lasting effect of warming and nitrogen addition. It was found that IDD determined the decomposition dynamics of litter during the recovery period, which masked the contribution of SST and SSW to litter decomposition, which may be related to the priming effect of initial soil environmental factors (Liu et al. 2023). In this study, the beginning of the treatment period was January, while the beginning of the recovery period was April (Fig. 3a), and the decomposition rate of the first decomposition stage was significantly higher in the recovery period than in the treatment period, similar to our previous study (Liu et al. 2023), which also changed the earlier IDD and thus the whole decomposition stage. Most importantly, the recovery period results confirmed the treatment effects of warming and nitrogen addition in the treatment period.

In summary, warming and nitrogen addition inhibited litter decomposition in subtropical forests, but there was no interaction between them. Warming and nitrogen addition significantly reduced the litter decomposition rate by 35.8% and 23.0%, respectively. Variance partitioning analysis (VPA) showed that stage soil temperature (SST) and soil water content (SSW) had positive effects on the stage decomposition rate (SDR), and the contribution rate of SST (35.5%) was greater than that of SSW (17.4%). The recovery experiment confirmed the effect of warming and nitrogen addition. During the recovery period, the litter decomposition rate recovered, confirming the effects of the treatments on litter decomposition. The results of this study further clarified the impact of nitrogen deposition on litter decomposition in subtropical forests under the background of climate warming.

Acknowledgement

We thank Qi Luo and Wen-Zheng Yang for their assistance with the field experiment. We thank the management authority of the Ailao Mountains National Nature Reserve and Ailaoshan Station for Subtropical Forest Ecosystem Studies for granting us permissions and facilitating this research. We are also grateful to the anonymous referees for their valuable comments, constructive suggestions and editorial assistance, which have greatly improved the paper. This study was supported by the Natural Science Foundation of Anhui Province (2108085QC101), Natural Science Foundation of Universities of Anhui Province for Distinguished Young Project (2022AH020081), and Natural Science Foundation of Universities of Anhui Province (No. KJ2020A0079). Opening Project of Anhui Province Key Laboratory of Environmental Hormone and Reproduction, Fuyang Normal University (No. FSKFKT012).

Author contributions

Yue Li presented the main idea and finished the initial draft. Shuai Liu and Changsheng Wu extensively modified the whole manuscript. Material preparation and data collection were performed by Simeng Wang, Ying Wang, and Menglong Bu. All authors read and approved the final manuscript.

Data availability

Data generated or analyzed during this study are available from the corresponding author upon reasonable request.

Competing interests

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

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published 11 Apr, 2024

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Editorial decision: Major revisions
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Climate warming and nitrogen deposition had no significant interaction effect on litter decomposition of subtropical mountain forest in Southwest China

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Journal Publication

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Abstract

Background and Aims

Methods

Results

Conclusions

Figures

1. Introduction

2. Materials and methods

2.1 Study site description

2.2 Experimental design and setup

2.3 Data collection and analysis

3. Results

3.1 Effects on soil microenvironmental factors

3.2 Effects of warming and nitrogen addition on litter decomposition

3.3 Factors affecting litter decomposition

4. Discussion

5. Conclusions

Declarations

References

Status:

Journal Publication

Version 1