文献信息:Widdig Meike, Heintz-Buschart Anna, Schleuss Per-Marten, Guhr Alexander, Borer Elizabeth T, Seabloom Eric W, Spohn Marie. Effects of nitrogen and phosphorus addition on microbial community composition and element cycling in a grassland soil. Soil Biology and Biochemistry, 2020, 108041. DOI: 10.1016/j.soilbio.2020.108041
摘 要:
微生物介导了土壤中的养分循环,因此普遍认为微生物在很大程度上控制着陆地生态系统对人为养分输入的响应。因此,有必要了解氮(N)和磷(P)有效性的增加是如何影响土壤原核生物和真菌群落组成,以及群落组成的变化如何影响元素循环。本研究通过温带草原一个9年的氮磷添加试验,测定了土壤微生物群落和土壤元素循环过程。
氮添加会影响微生物群落的组成,而原核生物群落对氮的敏感性不如真菌群落。
磷添加对微生物群落组成的影响较小,这表明该草原上磷对于微生物类群来说,选择性不如氮。
土壤pH和总有机碳(C)含量是与原核生物群落组成有关的主要因素,而可溶性有机碳与可溶性性氮的比例是真菌群落组成的主要驱动力。
这与本研究预先假设相反,植物生物量和植物群落结构仅解释了微生物群落组成的一小部分。
尽管微生物群落组成随养分添加而变化,但微生物生物量含量和呼吸速率没有改变
,这表明
微生物群落功能冗余
。
与植物生物量、植物群落结构或微生物群落相比,非生物因素对微生物呼吸作用、净氮矿化和非共生固氮作用的调控更强
,这表明
在养分输入增加的情况下,群落的转变不一定反映在元素循环速率上
。本研究表明,
大气氮沉降对真菌组成的影响可能比对原核生物的影响更大,且养分输入直接影响元素循环速率,而不是通过植物或微生物群落组成的变化来介导。
Graphical abstract
Table 1.
Soil pH, total organic carbon (TOC), total nitrogen (TN), and total phosphorus (TP) contents, dissolved organic carbon (DOC), dissolved nitrogen (DN), and dissolved inorganic phosphorus (DIP) concentrations under N and P addition in 0–15 cm and 15–30 cm soil depth.
Numbers depict means ± standard deviations (n = 3). Two-way ANOVA was conducted followed by Tukey test for multiple comparisons. Lower-case letters indicate significant differences between treatments tested separately for each depth increment. If no lower-case letters are shown, treatments did not differ significantly. Asterisks indicate significant differences between depth increments tested individually for each treatment.
1 Data were log 10 transformed for statistical tests,
2 reciprocally transformed (1/x) for statistical tests.
Table 2.
Microbial biomass carbon (MBC), microbial respiration, microbial respiration per unit MBC (qCO2), net nitrogen (N) mineralization, and non-symbiotic nitrogen (N2) fixation under N and P addition in 0–15 cm and 15–30 cm soil depth.
Numbers depict means ± standard deviations (n = 3). Two-way ANOVA was conducted followed by Tukey test for multiple comparisons. Lower-case letters indicate significant differences between treatments tested separately for each depth increment. If no lower-case letters are shown, treatments did not differ significantly. Asterisks indicate significant differences between depth increments tested individually for each treatment.
Table 3.
Multiple regression analysis after backward stepwise selection for identification of environmental controls on microbial respiration, net nitrogen (N) mineralization, and non-symbiotic N2 fixation in 0–15 cm depth.
The initial model contained soil pH, total organic carbon (TOC), total N (TN) contents, dissolved organic carbon (DOC), and dissolved N (DN) concentrations, DOC:DN ratio, prokaryotic and fungal community composition (based on first axis of principal coordinates analysis), and plant biomass and diversity. Displayed is the first model with a p-value below 0.05 and the highest number of remaining variables to show the influence of several variables. Variance inflation factors were used to check for multicollinearity and highly collinear variables were dropped.
Fig. 1.
Proportion of microbial respiration (
a
), net nitrogen (N) mineralization (
b
), and non-symbiotic N2 fixation rates (
c
) in 0–15 cm soil depth explained by the displayed soil, microbial, and plant factors.
Soil factors include soil pH, total organic carbon (TOC), dissolved organic carbon-to-dissolved nitrogen ratio (DOC:DN), total nitrogen (TN), dissolved organic carbon (DOC), and dissolved nitrogen (DN). Microbial factors include prokaryotic and fungal community composition at ASV level based on first axis of principal coordinates analysis. Plant factors include plant biomass, and plant diversity measured as Shannon diversity. All input variables are displayed, for a significant model, variables were removed stepwise as displayed in Table 3.
Fig. 2.
Prokaryotic (
a
) and fungal (
b
) community composition at ASV level displayed via non-metric multidimensional scaling (NMDS) of Jensen-Shannon divergences for different treatments in 0–15 cm soil depth.
To test for significant differences in community profiles of all treatments, analyses of similarity were performed on JSDs of both depth increments (Table S1) stratifying for sampling depth.
Fig. 3.
Proportion of variation (R2) of prokaryotic (
a
) and fungal (
b
) community composition explained by the displayed soil and plant factors in 0–15 cm soil depth.
Results are based on permutational multivariate analyses of variance (PERMANOVA) using Jensen-Shannon divergence of microbial communities at ASV level (Table S3), included are all factors with significant explanatory value in single-factor PERMANOVA. Soil factors include soil pH, total organic carbon (TOC), dissolved organic carbon-to-dissolved nitrogen ratio (DOC:DN), total nitrogen (TN), dissolved organic carbon (DOC), and dissolved nitrogen (DN). Plant factors include plant community based on the first axis of principal coordinates analysis, plant biomass, and plant diversity measured as Shannon diversity.
Fig. 4.
Relative abundances of prokaryotic genera in 0–15 cm soil depth.
Displayed are prokaryotic genera that made up > 2% of relative abundance. Prokaryotic genera <2% relative abundance were grouped as “Other”. Differentially abundant genera were detected from a data matrix containing the samples from the Ctrl, N10, N10P and P treatments with reads summed up at genus level, using DESeq2 with the model Y ∼ N * P. Differentially abundant genera are indicated with an asterisk in the legend and displayed in Table S4.
Fig. 5.
Relative abundances of fungal genera in 0–15 cm soil depth.
Displayed are genera with >1% relative abundance. Unclassified fungi and fungal genera <1% abundance were grouped as “Other”. Differentially abundant genera were detected from a data matrix containing the samples from the Ctrl, N10, N10P and P treatments with reads summed up at genus level, using DESeq2 with the model Y ∼ N * P. Differentially abundant genera are indicated with an asterisk in the legend and displayed in Table S4.