The use of fire as a tool to manipulate the environment has been instrumental in the human conquest of Earth, the first evidence of the use of fires by early hominids dating back to 1–1.5 million years ago [1]. Even today, most human-ignited vegetation fires take place on the African continent, and its widespread, frequently burned savannas bear ample witness to this. Although natural fires can occur even in tropical forest regions [2, 3], the extent of fires has greatly expanded on all continents with the arrival of Homo sapiens. Measurements of charcoal in dated sediment cores have shown clear correlations between the rate of burning and human settlement [4]. Pollen records show a shift with human settlement from pyrophobic vegetation to pyrotolerant and pyrophilic species, testimony to the large ecological impact of human-induced fires.

https://science.sciencemag.org/content/sci/250/4988/1669.full.pdf

Biomass Burning in the Tropics: Impact on Atmospheric Chemistry and Biogeochemical Cycles

Nutrient limitation to primary productivity and other biological processes is widespread in terrestrial ecosystems, and nitrogen (N) and phosphorus (P) are the most common limiting elements, both individually and in combination. Mechanisms that drive P limitation, and their interactions with the N cycle, have received less attention than mechanisms causing N limitation. We identify and discuss six mechanisms that could drive P limitation in terrestrial ecosystems. The best known of these is depletion-driven limitation, in which accumulated P losses during long-term soil and ecosystem development contribute to what Walker and Syers termed a "terminal steady state" of profound P depletion and limitation. The other mechanisms are soil barriers that prevent access to P; transactional limitation, in which weathering of P-containing minerals does not keep pace with the supply of other resources; low-P parent materials; P sinks; and anthropogenic changes that increase the supply of other resources (often N) relative to P. We distinguish proximate nutrient limitation (which occurs where additions of a nutrient stimulate biological processes, especially productivity) from ultimate nutrient limitation (where additions of a nutrient can transform ecosystems). Of the mechanisms that drive P limitation, we suggest that depletion, soil barriers, and low-P parent material often cause ultimate limitation because they control the ecosystem mass balance of P. Similarly, demand-independent losses and constraints to N fixation can control the ecosystem-level mass balance of N and cause it to be an ultimate limiting nutrient.

Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen–phosphorus interactions

Contents

 Summary

I Introduction
II Variability of N : P ratios in response to nutrient  supply
III Critical N : P ratios as indicators of nutrient  limitation
IV Interspecific variation in N : P ratios
V Vegetation properties in relation to N : P ratios
VI Implications of N : P ratios for human impacts  on ecosystems
VII Conclusions

 Acknowledgements


 References



Summary
Nitrogen (N) and phosphorus (P) availability limit plant growth in most terrestrial ecosystems. This review examines how variation in the relative availability of N and P, as reflected by N : P ratios of plant biomass, influences vegetation composition and functioning. Plastic responses of plants to N and P supply cause up to 50-fold variation in biomass N : P ratios, associated with differences in root allocation, nutrient uptake, biomass turnover and reproductive output. Optimal N : P ratios – those of plants whose growth is equally limited by N and P – depend on species, growth rate, plant age and plant parts. At vegetation level, N : P ratios 20 often (not always) correspond to N- and P-limited biomass production, as shown by short-term fertilization experiments; however long-term effects of fertilization or effects on individual species can be different. N : P ratios are on average higher in graminoids than in forbs, and in stress-tolerant species compared with ruderals; they correlate negatively with the maximal relative growth rates of species and with their N-indicator values. At vegetation level, N : P ratios often correlate negatively with biomass production; high N : P ratios promote graminoids and stress tolerators relative to other species, whereas relationships with species richness are not consistent. N : P ratios are influenced by global change, increased atmospheric N deposition, and conservation managment.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2004.01192.x

N : P ratios in terrestrial plants: variation and functional significance

The terrestrial biosphere plays an important role in the global carbon cycle. In the 1994 Intergovernmental Panel Assessment on Climate Change (IPCC), an effort was made to improve the quantification of terrestrial exchanges and potential feedbacks from climate, changing CO2, and other factors; this paper presents the key results from that assessment, together with expanded discussion. The carbon cycle is the fluxes of carbon among four main reservoirs: fossil carbon, the atmosphere, the oceans, and the terrestrial biosphere. Emissions of fossil carbon during the 1980s averaged 5.5 Gt y−1. During the same period, the atmosphere gained 3.2 Gt C y−1 and the oceans are believed to have absorbed 2.0 Gt C y−1. The regrowing forests of the Northern Hemisphere may have absorbed 0.5 Gt C y−1 during this period. Meanwhile, tropical deforestation is thought to have released an average 1.6 Gt C y−1 over the 1980s. While the fluxes among the four pools should balance, the average 198Ds values lead to a ‘missing sink’ of 1.4 Gt C y−1 Several processes, including forest regrowth, CO2 fertilization of plant growth (c. 1.0 Gt C y−1), N deposition (c. 0.6 Gt C y−1), and their interactions, may account for the budget imbalance. However, it remains difficult to quantify the influences of these separate but interactive processes. Uncertainties in the individual numbers are large, and are themselves poorly quantified. This paper presents detail beyond the IPCC assessment on procedures used to approximate the flux uncertainties.

Lack of knowledge about positive and negative feedbacks from the biosphere is a major limiting factor to credible simulations of future atmospheric CO2 concentrations. Analyses of the atmospheric gradients of CO2 and 13 CO2 concentrations provide increasingly strong evidence for terrestrial sinks, potentially distributed between Northern Hemisphere and tropical regions, but conclusive detection in direct biomass and soil measurements remains elusive.

Current regional-to-global terrestrial ecosystem models with coupled carbon and nitrogen cycles represent the effects of CO2 fertilization differently, but all suggest longterm responses to CO2 that are substantially smaller than potential leaf- or laboratory whole plant-level responses. Analyses of emissions and biogeochemical fluxes consistent with eventual stabilization of atmospheric CO2 concentrations are sensitive to the way in which biospheric feedbacks are modeled by c. 15%. Decisions about land use can have effects of 100s of Gt C over the next few centuries, with similarly significant effects on the atmosphere.

Critical areas for future research are continued measurements and analyses of atmospheric data (CO2 and 13CO2) to serve as large-scale constraints, process studies of the scaling from the photosynthetic response to CO2 to whole-ecosystem carbon storage, and rigorous quantification of the effects of changing land use on carbon storage.

Terrestrial ecosystems and the carbon cycle

Human activities have clearly caused dramatic alterations of the terrestrial nitrogen cycle, and analyses of the extent and effects of such changes are now common in the scientific literature. However, any attempt to evaluate N cycling processes within ecosystems, as well as anthropogenic influences on the N cycle, requires an understanding of the magnitude of inputs via biological nitrogen fixation (BNF). Although there have been many studies addressing the microbiology, physiology, and magnitude of N fixation at local scales, there are very few estimates of BNF over large scales. We utilized >100 preexisting published estimates of BNF to generate biome- and global-level estimates of biological N fixation. We also used net primary productivity (NPP) and evapotranspiration (ET) estimates from the Century terrestrial ecosystem model to examine global relationships between these variables and BNF as well as to compare observed and Century-modeled BNF. Our data-based estimates showed a strong positive relationship between ecosystem ET and BNF, and our analyses suggest that while the model's simple relationships for BNF predict broad scale patterns, they do not capture much of the variability or magnitude of published rates. Patterns of BNF were also similar to patterns of ecosystem NPP. Our “best estimate” of potential nitrogen fixation by natural ecosystems is ∼195 Tg N yr−1, with a range of 100–290 Tg N yr−1. Although these estimates do not account for the decrease in natural N fixation due to cultivation, this would not dramatically alter our estimate, as the greatest reductions in area have occurred in systems characterized by relatively low rates of N fixation (e.g., grasslands). Although our estimate of BNF in natural ecosystems is similar to previously published estimates of terrestrial BNF, we believe that this study provides a more documented, constrained estimate of this important flux.

/pdf/global-patterns-of-terrestrial-biological-nitrogen-n2-4j06hwerc7.pdf

Global patterns of terrestrial biological nitrogen (N2) fixation in natural ecosystems

Prescribed burning is a major control over element cycles in Tallgrass prairie (Eastern Kansas, USA) In this paper we report potential effects of fire on nonsymbiotic nitrogen fixation Fire resulted in additions of available P in ash, which may stimulate nitrogen fixation by terrestrial cyanobacteria Cyanobacterial nitrogenase activity and biomass responded positively to additions of ash or P in laboratory assays using soil Further assays in soil showed that cyanobacteria responded to changes in available N:available P ratio (aN:P) across a range of concentrations Nitrogen fixation rate could be related empirically to aN:P via a log-linear relationship Extrapolation of laboratory results to the field yielded a maximal estimate of 21 kg N ha-1 y-1 Results support arguments from the marine and terrestrial literature that P availability is central to regulation of ecosystem N budgets

K. A. Eisele

Papers

Effects of available P and N:P ratios on non-symbiotic dinitrogen fixation in tallgrass prairie soils