Nitrogen and phosphorus enrichment differentially affect grassland ecosystem functioning via multi-trophic pathways
Data files
Jul 02, 2025 version files 197.88 KB
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NP_Data.xlsx
94.25 KB
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R_code.txt
97.25 KB
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README.md
6.38 KB
May 08, 2026 version files 198.84 KB
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NP_Data_New.xlsx
93.93 KB
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R_code.txt
97.25 KB
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README.md
7.65 KB
Abstract
The enrichment of nutrients (e.g., N and P) is crucial in shaping the links between diversity and ecosystem functions. Previous research has mainly focused on how nutrient enrichment affects community diversity and composition within a single trophic level, such as plants or soil microbial communities. However, how nutrient enrichment affects community composition and diversity across multiple trophic levels and ecosystem functions has not been extensively explored. By conducting two long-term N- and P-enrichment experiments in temperate steppes, we explored the response of diversity and composition at both the community and functional group level across the soil food webs to nutrient enrichment, as well as their relationship with ecosystem functions. We found that the responses of soil biota and plants to nutrient enrichment varied depending on the community and functional group. N-enrichment resulted in a decrease in plant and bacterial richness at both the community and functional group levels, while P-enrichment decreased fungal and nematode richness at both levels. In addition, N-enrichment changed the community composition of plants and all soil biota, while P-enrichment only affected the community composition of fungi and nematodes. Our findings also showed that community composition across multiple trophic levels played a crucial role in regulating ecosystem functions (e.g., plant biomass, soil biota biomass, and soil organic matter (SOM) decomposition) when considering both community composition and diversity simultaneously. For example, N-enrichment induced changes in the community composition of plants, bacteria, and nematodes, which altered plant biomass, while P-enrichment induced changes in the community composition of nematodes that altered soil biota biomass and SOM decomposition. Overall, our study suggests that long-term nutrient enrichment may have a greater impact on community composition than diversity across multiple trophic levels, and these changes in community composition can have broad implications for ecosystem functions. Therefore, more effort should be made not only on community diversity but also on composition to clarify the relationship between diversity and ecosystem functions in future studies under climate change.
The dataset consists of an NP_Data_New.xlsx.xlsx and an R code file (R_code.txt). Sheet 1 was the dataset for all raw data. The dataset contains 113 variables for Sheet 1. NA in the dataset indicates missing data for the relevant variable.
Column name -- Units, Description
Column name -- Units, Description
SampleID -- Unique ID for each sample
Treat -- unitless Different treatments
Block -- Unique ID for each block
Lev -- unitless Level of nutrient enrichment
ZLev1 -- unitless Level of nutrient enrichment
ZLev -- unitless Level of nutrient enrichment
PH -- Soil pH, Natural log transformation
SM -- % Soil moisture, Natural log transformation
TIN -- mg kg-1 Total inorganic nitrogen, Natural log transformation
SOC -- g kg-1 Soil organic carbon, Natural log transformation
TSN -- g kg-1 Total soil nitrogen, Natural log transformation
TSP -- g kg-1 Total soil phosphorus, Natural log transformation
CTN -- ratio of SOC and TSN, Natural log transformation
CTP -- ratio of SOC and TSP, Natural log transformation
NTP -- ratio of TSN and TSP, Natural log transformation
Plant_Rich -- Plant richness, Natural log transformation
AB_Rich -- Annuals and biennials richness, Natural log transformation
PB_Rich -- Perennial bunchgrass richness, Natural log transformation
PF_Rich -- Perennial forbs richness, Natural log transformation
PR_Rich -- Perennial rhizome grasses richness, Natural log transformation
Ba_Rich -- Bacterial richness, Natural log transformation
BC_Rich -- C-cycle bacteria richness, Natural log transformation
BN_Rich -- N-cycle bacteria richness, Natural log transformation
Fu_Rich -- Fungal richness, Natural log transformation
ECM_Rich -- Mycorrhizal fungi richness, Natural log transformation
Path_Rich -- Plant pathogen fungi richness, Natural log transformation
Sap_Rich -- Soil saprotrophic fungi composition, Natural log transformation
Nema_Rich -- Soil nematode richness, Natural log transformation
BF_Rich -- Bacteria-feeders nematode richness, Natural log transformation
FF_Rich -- Fungal-feeders nematode richness, Natural log transformation
HF_Rich -- Plant-feeders nematode richness, Natural log transformation
OC_Rich -- Omnivores+carnivores nematode richness, Natural log transformation
Plant_PCo1 -- Plant composition, first axis of the PCoA
Plant_PCo2 -- Plant composition, second axis of the PCoA
AB_PCo1 -- Annuals and biennials composition, first axis of the PCoA
AB_PCo2 -- Annuals and biennials composition, second axis of the PCoA
PB_PCo1 -- Perennial bunchgrass composition, first axis of the PCoA
PB_PCo2 -- Perennial bunchgrass composition, second axis of the PCoA
PF_PCo1 -- Perennial forbs composition, first axis of the PCoA
PF_PCo2 -- Perennial forbs composition, second axis of the PCoA
PR_PCo1 -- Perennial rhizome grasses composition, first axis of the PCoA
PR_PCo2 -- Perennial rhizome grasses composition, second axis of the PCoA
Ba_PCo1 -- Bacterial composition, first axis of the PCoA
Ba_PCo2 -- Bacterial composition, second axis of the PCoA
BC_PCo1 -- C-cycle bacteria composition, first axis of the PCoA
BC_PCo2 -- C-cycle bacteria composition, second axis of the PCoA
BN_PCo1 -- N-cycle bacteria composition, first axis of the PCoA
BN_PCo2 -- N-cycle bacteria composition, second axis of the PCoA
Fu_PCo1 -- Fungal composition, first axis of the PCoA
Fu_PCo2 -- Fungal composition, second axis of the PCoA
ECM_PCo1 -- Mycorrhizal fungi composition, first axis of the PCoA
ECM_PCo2 -- Mycorrhizal fungi composition, second axis of the PCoA
Path_PCo1 -- Plant pathogen fungi composition, first axis of the PCoA
Path_PCo2 -- Plant pathogen fungi composition, second axis of the PCoA
Sap_PCo1 -- Soil saprotroph fungi composition, first axis of the PCoA
Sap_PCo2 -- Soil saprotroph fungi composition, second axis of the PCoA
Nema_PCo1 -- Nematode taxon composition, first axis of the PCoA
Nema_PCo2 -- Nematode taxon composition, second axis of the PCoA
BF_PCo1 -- Bacterial feeders composition, first axis of the PCoA
BF_PCo2 -- Bacterial feeders composition, second axis of the PCoA
FF_PCo1 -- Fungal feeders composition, first axis of the PCoA
FF_PCo2 -- Fungal feeders composition, second axis of the PCoA
HF_PCo1 -- Plant feeders composition, first axis of the PCoA
HF_PCo2 -- Plant feeders composition, second axis of the PCoA
OC_PCo1 -- Omnivores+carnivores composition, first axis of the PCoA
OC_PCo2 -- Omnivores+carnivores composition, second axis of the PCoA
ZPlant_Rich -- Min-max normalized plant richness
ZPlant_PCoA1 -- Min-max normalized plant composition
ZAB_Rich -- Min-max normalized annuals and biennials richness
ZAB_PCoA1 -- Min-max normalized annuals and biennials composition
ZPB_Rich -- Min-max normalized perennial bunchgrass richness
ZPB_PCoA1 -- Min-max normalized perennial bunchgrass composition
ZPF_Rich -- Min-max normalized perennial forbs richness
ZPF_PCoA1 -- Min-max normalized perennial forbs composition
ZPR_Rich -- Min-max normalized perennial rhizome grasses richness
ZPR_PCoA1 -- Min-max normalized perennial rhizome grasses composition
ZBa_Rich -- Min-max normalized bacterial richness
ZBa_PCoA1 -- Min-max normalized bacterial composition
ZBC_Rich -- Min-max normalized C-cycle bacteria richness
ZBC_PCoA1 -- Min-max normalized C-cycle bacteria composition
ZBN_Rich -- Min-max normalized N-cycle bacteria richness
ZBN_PCoA1 -- Min-max normalized N-cycle bacteria composition
ZFu_Rich -- Min-max normalized fungal richness
ZFu_PCoA1 -- Min-max normalized fungal composition
ZECM_Rich -- Min-max normalized mycorrhizal fungi richness
ZECM_PCoA1 -- Min-max normalized mycorrhizal fungi composition
ZPath_Rich -- Min-max normalized plant pathogen fungi richness
ZPath_PCoA1 -- Min-max normalized plant pathogen fungi composition
ZSap_Rich -- Min-max normalized soil saprotroph fungi richness
ZSap_PCoA1 -- Min-max normalized soil saprotroph fungi composition
ZNema_Rich -- Min-max normalized soil nematode richness
ZNema_PCoA1 -- Min-max normalized nematode taxon composition
ZBF_Rich -- Min-max normalized bacteria-feeders nematode richness
ZBF_PCoA1 -- Min-max normalized bacterial feeders composition
ZFF_Rich -- Min-max normalized fungal-feeders nematode richness
ZFF_PCoA1 -- Min-max normalized fungal feeders composition
ZHF_Rich -- Min-max normalized plant-feeders nematode richness
ZHF_PCoA1 -- Min-max normalized plant feeders composition
ZOC_Rich -- Min-max normalized omnivores+carnivores nematode richness
ZOC_PCoA1 -- Min-max normalized omnivores+carnivores composition
AFun -- the functions of plant biomass, Natural log transformation
BFun -- the functions of soil biota biomass, Natural log transformation
SFun -- the functions of SOM decomposition, Natural log transformation
Litter -- Soil decomposition, Natural log transformation
ANPP -- g m-1 Aboveground net primary production, Natural log transformation
BB -- g m-1 Belowground biomass, Natural log transformation
Cmin -- mg kg-1 day-1 Soil C mineralization rate, Natural log transformation
Nmin -- mg kg-1 day-1 Soil N mineralization rate, Natural log transformation
TMB -- nmol g-1 Total microbial biomass, Natural log transformation
TNA -- ind g-1 Soil nematode abundance, Natural log transformation
ZAFun -- Min-max normalized the functions of plant biomass
ZBFun -- Min-max normalized the functions of soil biota biomass
ZSFun -- Min-max normalized the functions of SOM decomposition
Code/Software
The file R_code includes all the code required for the manuscript.
Any scripts, code,or packages can be run using R version 4.0.5 (R Core Team, 2021).
after Jul 2, 2025: A discrepancy was caused by a data compilation error during the preparation of the original Excel file. Specifically, when merging data from multiple source sheets, the columns containing PCoA values (e.g., Plant_PCo1, Plant_PCo2, AB_PCo1, AB_PCo2, and all related PCoA columns up to OC_PCo2) were inadvertently pasted out of order relative to the SampleIDs. As a result, the PCoA values in the original file do not correctly correspond to their respective samples.
Importantly, this error does not affect the results or conclusions presented in our published article (Journal of Ecology, 2025, 113: 2447–2459). In all statistical analyses reported in the paper, we used min‑max normalized data (e.g., ZPlant_Rich, ZPlant_PCoA1, ZAB_Rich, ZAB_PCoA1, etc.), which are correctly calculated and presented in the original file (columns BO to CF). These min‑max normalized values were derived from the correct richness and PCoA data and are accurately aligned. Detailed data transformation procedures can be found in the README.md file. All figures and tables in the article are based on these min‑max normalized variables, so the integrity of the published work remains fully intact.
We sincerely apologize for this clerical error in the dataset preparation and for any confusion it may have caused. We have now uploaded the corrected Excel file and README.md file.
