globalchange  > 气候变化事实与影响
DOI: doi:10.1038/nclimate2549
论文题名:
Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes
作者: Z. Y. Yuan
刊名: Nature Climate Change
ISSN: 1758-992X
EISSN: 1758-7112
出版年: 2015-03-09
卷: Volume:5, 页码:Pages:465;469 (2015)
语种: 英语
英文关键词: Ecosystem ecology ; Environmental monitoring
英文摘要:

Living organisms maintain a balance of chemical elements for optimal growth and reproduction, which plays an important role in global biogeochemical cycles1, 2, 3, 4, 5. Human domination of Earth’s ecosystems has led to drastic global changes6, 7, 8, but it is unclear how these affect the stoichiometric coupling of nutrients in terrestrial plants, the most important food source on Earth. Here we use meta-analyses of 1,418 published studies to show that the ratio of terrestrial plant nitrogen (N) to phosphorus (P) decreases with elevated concentrations of CO2, increasing rainfall, and P fertilization, but increases with warming, drought, and N fertilization. Our analyses also reveal that multiple global change treatments generally result in overall additive effects of single-factor treatments and that the responses of plant nutrients and their stoichiometry are similar in direction, but often greater in controlled than in natural environments. Our results suggest a decoupling of the P biogeochemical cycle from N in terrestrial plants under global changes6, 7, 8, which in turn may diminish the provision of ecosystem services1, 5, 9.

From cellular metabolism to ecosystem structure and nutrient cycling, C, N and P are biologically coupled through their effects on the biochemical reactions that control primary production, respiration and decomposition in terrestrial ecosystems1, 2, 3, 4, 5, 8, 10, 11, 12. In the biosphere, living organisms, the major part of biogeochemical cycles, require elements in strict proportions to catalyse metabolic reactions and synthesize essential compounds with specific ratios of C:N:P (refs 1, 2). The biological control—that is, the conserved elemental stoichiometry of organisms—couples biogeochemical cycles to one another3. However, owing to different degrees of control by biological and geochemical processes, biogeochemical C, N and P cycles could be unbalanced or decoupled under rapid global changes2, 8, 13. For example, an increase in aridity with climate changes can reduce soil C and N, but increase soil P in global drylands13, indicating that the coupling between biogeochemical cycles is fragile in drylands in the face of rapid climate change. The decoupling of the biogeochemical cycles of C, N and P may also lead to nutrient decoupling in plants that form the base of food chains5, 8 and consequently can negatively influence the trophic structures and the services of terrestrial ecosystems14.

Global changes have drastically affected the biogeochemical cycles of carbon and nutrient elements of Earth’s ecosystems6, 15. The simultaneous changes in global-scale biogeochemical cycles (for example, elevated CO2 concentration [CO2], atmospheric N deposition, and N and P fertilization) and in climates (increasing temperature and altered rainfall) are anticipated to have stoichiometric consequences worldwide (Fig. 1). For example, elevated [CO2] can increase plant C fixation, but stimulated plant photosynthesis, growth and overall production may lead to decreases of plant nutrient concentrations—that is, the ‘dilution effect’16, 17. Warming tends to increase soil microbial activity, but may induce warming-associated droughts, both of which affect plant photosynthesis and plant stoichiometry. The same is true for changes in precipitation that affect plant stoichiometry via soil water availability.

Figure 1: A conceptual diagram of the influence of global changes on processes controlling the stoichiometry of plant C, N and P.
A conceptual diagram of the influence of global changes on processes controlling the stoichiometry of plant C, N and P.

Rectangles are nutrient pools, hexagons indicate biogeochemical processes and valves (red symbols) are controls on plant C, N and P. Plant illustration © Elena Belyakova/Thinkstock.

We searched databases of ISI Web of Science, PubMed, Google Scholar and JSTOR (Supplementary References). Our data covered a wide range of all terrestrial ecosystem types including Arctic tundra, forests, and grasslands (Supplementary Fig. 1). All original data were extracted from the text, tables, figures and appendices in the publications. When data were presented graphically, numerical data were obtained by Image-Pro Plus 7.0 (Media Cybernetics). Measurements from different ecosystem types, species, plant tissue types and treatment levels within a single study were considered independent observations. If multiple observations from different sampling dates at the same site were reported, we used the first observation in the analysis (Supplementary Data 1).

To examine the effects of global change treatments on plant stoichiometric C, N and P, we calculated response ratios from each individual study as described in ref. 30. Natural log response ratio (lnRR) was calculated as ln(Xe/Xc) = lnXe − lnXc, where Xe and Xc are the response values of each individual observation in the treatment and in the control, respectively. The corresponding sampling variance for each lnRR was calculated as ln[(1/ne) × (Se/Xe)2 + (1/nc) × (Sc/Xc)2] in R with the package ‘metafor’ 1.9-2, where ne, nc, Se, Sc, Xe and Xc are sample sizes, standard deviations and mean response values in the treatment and in the control, respectively. The natural log response ratios to individual and combined treatments were determined by specifying studies as random factor using the rma model in metafor. The effects of global change treatments on plant stoichiometric C, N and P were considered significant if the 95% confidence intervals (CI) of lnRR did not overlap zero. To compare whether the responses of plant nutrients and their stoichiometric ratios differ between studies conducted in natural environments and controlled environments, such as greenhouse and growth chamber experiments, we compared estimated lnRR by their CIs.

To examine whether treatment effects are additive on plant nutrients and their stoichiometric ratios, we tested whether the interactions between multiple treatments are significant by using rma.uni models in metafor with treatments as categorical predictors. A significant interaction between treatments indicates that the treatment effects are not additive. Because of limited data for three or more combined treatments, we considered only two-way interactions. In addition, we performed paired meta-analyses31, a more conservative comparison in which interactive effects of all observations from multiple-factor studies were examined by comparing the sum of effect sizes of single factors with the effect size of combined factors. Individual experiments of synergistic, antagonistic and additive responses should be situated above, below and across the 1:1 line, respectively31.

Because global change treatments vary strongly in quantities applied within and among studies, we examined the sensitivities of plant nutrients and their stoichiometric ratios to the quantities applied for all global change treatments by using REML estimation in the rma.uni model for metafor with the applied rates of global change treatments as continuous variables. In this analysis, we considered only the responses of plant nutrients and their stoichiometric ratios to individual global change treatments owing to limited data availability for multiple treatments as well as additivity of multiple treatments. To examine whether N:P responded to water addition, or drought experiments differ with background water availability, we derived an aridity index, given by the ratio of precipitation to potential evapotranspiration, by using data interpolations provided by WorldClim (http://WorldClim.org) and by CGIAR-CSI (http://www.cgiar-csi.org/data/global-aridity-and-pet-database). We then tested whether the log N:P response ratio changed with the aridity index. All statistical analyses were performed in R 3.0.2.

  1. Sterner, R. W. & Elser, J. J. Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere (Princeton Univ. Press, 2002).
  2. Finzi, A. C. et al. Responses and feedbacks of coupled biogeochemical cycles to climate change: Examples from terrestrial ecosystems. Front. Ecol. Environ. 9, 6167 (2011).
  3. Schlesinger, W. H., Cole, J. J., Finzi, A. C. & Holland, E. A. Introduction to coupled biogeochemical cycles. Front. Ecol. Environ. 9, 58 (2011).
  4. Rivas-Ubach, A., Sardans, J., Perez-Trujillo, M., Estiarte, M. & Peñuelas, J. Strong relationship between elemental stoichiometry and metabolome in plants. Proc. Natl Acad. Sci. USA 109, 41814186 (2012).
  5. Peñuelas, J. et al. Human-induced nitrogen–phosphorus imbalances alter natural and managed ecosystems across the globe. Nature Commun. 4, 2934 (2013).
  6. Vitousek, P. M., Mooney, H. A., Lubchenco, J. & Melillo, J. M. Human domination of Earth’s ecosystems. Science 277, 494499 (1997).
  7. Diffenbaugh, N. S. & Field, C. B. Changes in ecologically critical terrestrial climate conditions. Science 341, 486492 (2013).
  8. Peñuelas, J., Sardans, J., Rivas-Ubach, A. & Janssens, I. A. The human-induced imbalance between C, N and P in Earth’s life system. Glob. Change Biol. 18, 36 (2012).
  9. Elser, J. J. et al. Nutritional constraints in terrestrial and freshwater food webs. Nature 408, 578580 (2000).
URL: http://www.nature.com/nclimate/journal/v5/n5/full/nclimate2549.html
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资源类型: 期刊论文
标识符: http://119.78.100.158/handle/2HF3EXSE/4820
Appears in Collections:气候变化事实与影响
科学计划与规划
气候变化与战略

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Z. Y. Yuan. Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes[J]. Nature Climate Change,2015-03-09,Volume:5:Pages:465;469 (2015).
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