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

Humans now strongly influence almost every major aquatic ecosystem, and their activities have dramatically altered the fluxes of growth-limiting nutrients from the landscape to receiving waters. Unfortunately, these nutrient inputs have had profound negative effects upon the quality of surface waters worldwide. This review examines how eutrophication influences the biomass and species composition of algae in both freshwater and costal marine systems. An overview of recent advances in algae-related eutrophication research is presented. In freshwater systems, a summary is presented for lakes and reservoirs; streams and rivers; and wetlands. A brief summary is also presented for estuarine and coastal marine ecosystems. Eutrophication causes predictable increases in the biomass of algae in lakes and reservoirs; streams and rivers; wetlands; and coastal marine ecosystems. As in lakes, the response of suspended algae in large rivers to changes in nutrient loading may be hysteretic in some cases. The inhibitory effects of high concentrations of inorganic suspended solids on algal growth, which can be very evident in many reservoirs receiving high inputs of suspended soils, also potentially may occur in turbid rivers. Consistent and predictable eutrophication-caused increases in cyanobacterial dominance of phytoplankton have been reported worldwide for natural lakes, and similar trends are reported here both for phytoplankton in turbid reservoirs, and for suspended algae in a large river. A remarkable unity is evident in the global response of algal biomass to nitrogen and phosphorus availability in lakes and reservoirs; wetlands; streams and rivers; and coastal marine waters. The species composition of algal communities inhabiting the water column appears to respond similarly to nutrient loading, whether in lakes, reservoirs, or rivers. As is true of freshwater ecosystems, the recent literature suggests that coastal marine ecosystems will respond positively to nutrient loading control efforts. Our understanding of freshwater eutrophication and its effects on algal-related water quality is strong and is advancing rapidly. However, our understanding of the effects of eutrophication on estuarine and coastal marine ecosystems is much more limited, and this gap represents an important future research need. Although coastal systems can be hydrologically complex, the biomass of marine phytoplankton nonetheless appears to respond sensitively and predictably to changes in the external supplies of nitrogen and phosphorus. These responses suggest that efforts to manage nutrient inputs to the seas will result in significant improvements in coastal zone water quality. Additional new efforts should be made to develop models that quantitatively link ecosystem-level responses to nutrient loading in both freshwater and marine systems.

http://globalpnetwork.net/sites/default/files/Smith_2003_ESPR_eutrophication_review.pdf

Eutrophication of freshwater and coastal marine ecosystems a global problem

Nutrient limitation (mostly N or P) is a driving force in ecosystem development. Current techniques to determine the nature of nutrient limitation use laborious fertilization experiments. It was hypothesized that the N:P ratio of'the vegetation directly indicates the nature of nutrient limitation on a community level (N vs. P limitation). This hypothesis was tested by reviewing data on fertilization studies in a variety of European freshwater wetland ecosystems (bogs, fens, wet heathlands, dune slacks, wet grasslands). In a subset of the data (dune slacks) between-site intraspecific variation and within-site interspecific variation in nutrient content and N:P ratio was studied in five plant species. A review of 40 fertilization studies reveals that an N:P ratio >16 indicates P limitation on a community level, while an N:P ratio < 14 is indicative of N limitation. At N:P ratios between 14 and 16, either N or P can be limiting or plant growth is colimited by N and P together. In only one out of 40 fertilization studies, the N:P ratio gave a false indication of the nature of nutrient limitation. Measuring the N:P ratio of the vegetation is a simple and cheap alternative to fertilization studies. The method can only be used under conditions where either N or P controls plant growth. The dataset contains a large variety of vegetation types and plant species, and 11 I of the 40 sites were near-monocultures. This suggests that interspecific differences in critical N:P ratios among species may be insignificant. However, a rigorous test of this hypothesis is required. A survey in 18 dune slacks showed large within-site variation in N:P ratio among five species (Calamagrostis epigejos, Phragmites australis, Lycopus europaeus, Mentha aquatica and Eupatorium cannabinum). The N:P ratios of the five species suggested that within plant communities species can be differentially limited by N or P. Moreover, species with an N:P ratio that suggested P-limitation were found at sites where N controlled community biomass production, and vice versa. Between-site intraspecific variation in N and P contents and N:P ratios was also large, and about equal for the five species. This illustrates the plasticity of plant species with respect to N and P contents, probably in response to differences in N and P supply ratios. The vegetation N:P ratio is of diagnostic value and its use may increase our understanding of numerous facets of physiological, population, community and ecosystem ecology.

The vegetation N:P ratio: a new tool to detect the nature of nutrient limitation

Phytoplankton can become limited by the availability of nutrients when light and temperature are adequate and loss rates are not excessive. The current paradigms for nutrient limitations in freshwater, estuarine, and marine environments are quite different. A review of the experimental and observational data used to infer P or N limitation of phytoplankton growth indicates that P limitation in freshwater environments can be demonstrated rigorously at several hierarchical levels of system complexity, from algal cultures to whole lakes. A similarly rigorous demonstration of N limitation has not been achieved for marine waters. Therefore, we conclude that the extent and severity of N limitation in the marine environment remain an open question. Culture studies have established that internal cellular concentrations of nutrients determine phytoplankton growth rates, and these studies have shown that it is often difficult to relate growth rates to external concentrations, especially in natural situations. This should lead to a greater reliance on the composition of particulate matter and biomass-based physiological rates to infer nutrient limitation. Such measurements have demonstrated their utility in a wide variety of freshwater and marine environments, and, most importantly, they can be applied to systems that are difficult to manipulate experimentally or budget accurately. Dissolved nutrient concentrations are most useful in determining nutrient loading rates of aquatic ecosystems. The relative proportions of nutrients supplied to phytoplankton can be a strong selective force shaping phytoplankton communities and affecting the biomass yield per unit of limiting nutrient. A current dogma of aquatic science is that marine and estuarine phytoplankton tend to be nitrogen limited, while freshwater phytoplankton tend to be phosphorus limited. Carpenter and Capone (1983) documented the preeminence of N studies in the literature on brackish and marine ecosystems. In 1970 there were equal numbers of references per year to N and P. The decade of the seventies saw a nearly fourfold increase in references to N, while the number of P references per year remained essentially unchanged. No trend was evident for the freshwater literature despite the fact that by the late seventies the evidence for P limitation had become so great that phosphorus control was recommended as the legislated basis for controlling eutrophication in North American and European inland waters (e.g.

/pdf/nutrient-limitation-of-phytoplankton-in-freshwater-and-1gyy8avtuu.pdf

Nutrient limitation of phytoplankton in freshwater and marine environments: A review of recent evidence on the effects of enrichment1

Ecological stoichiometry is the study of the balance of multiple chemical elements in ecological interactions. This paper reviews recent findings in this area and seeks to broaden the stoichiometric concept for use in evolutionary studies, in integrating ecological dynamics with cellular and genetic mechanisms, and in developing a unified means for studying diverse organisms in diverse habitats. This broader approach would then be considered “biological stoichiometry”. Evidence supporting a hypothesised connection between the C:N:P stoichiometry of an organism and its growth rate (the “growth rate hypothesis”) is reviewed. Various data indicate that rapidly growing organisms commonly have low biomass C:P and N:P ratios. Evidence is then discussed suggesting that low C:P and N:P ratios in rapidly growing organisms reflect increased allocation to P-rich ribosomal RNA (rRNA), as rapid protein synthesis by ribosomes is required to support fast growth. Indeed, diverse organisms (bacteria, copepods, fishes, others) exhibit increased RNA levels when growing actively. This implies that evolutionary processes that generate, directly or indirectly, variation in a major life history trait (specific growth rate) have consequences for ecological dynamics due to their effects on organismal elemental composition. Genetic mechanisms by which organisms generate high RNA, high growth rate phenotypes are discussed next, focusing on the structure and organisation of the ribosomal RNA genes (the “rDNA”). In particular, published studies of a variety of taxa suggest an association between growth rate and variation in the length and content of the intergenic spacer (IGS) region of the rDNA tandem repeat unit. In particular, under conditions favouring increased growth or yield, the number of repeat units (“enhancers”) increases (and the IGS increases in length), and transcription rates of rRNA increase. In addition, there is evidence in the literature that increased numbers of copies of rDNA genes are associated with increased growth and production. Thus, a combination of genetic mechanisms may be responsible for establishing the growth potential, and thus the RNA allocation and C:N:P composition, of an organism. Furthermore, various processes, during both sexual and asexual reproduction, can generate variation in the rDNA to provide the raw material for selection and to generate ecologically significant variation in C:N:P stoichiometry. This leads us to hypothesize that the continuous generation of such variation may also play a role in how species interactions develop in ecosystems under different conditions of energy input and nutrient supply.

Biological stoichiometry from genes to ecosystems.

An analysis of mean growing season concentrations of chlorophyll, total phosphorus (TP), and total nitrogen (TN) in 228 north latitude lakes confirms previous observations that chlorophyll yield is dependent both on the phosphorus concentration and on the TN:TP ratio. Of two modified chlorophyll models which depend explicitly on both nitrogen and phosphorus developed and tested, one, a multiple regression model, appears to greatly reduce the error of chlorophyll prediction in lakes. A theoretical framework is presented which provides an explanation for the observed effects of N:P ratios.

https://aslopubs.onlinelibrary.wiley.com/doi/epdf/10.4319/lo.1982.27.6.1101

The nitrogen and phosphorus dependence of algal biomass in lakes: An empirical and theoretical analysis1

An analysis of growing season measurements of daily primary productivity, chlorophyll, water chemistry, and transparency from 58 north temperate lakes shows a strong correlation between volumetric rates of photosynthesis, chlorophyll, and nutrients. Mean daily rates of photosynthesis per unit volume euphotic zone, v, are correlated with mean chlorophyll concentration (r2 = 0.80). The mean daily rate of photosynthesis at optimal depth, Āopt, is highly correlated with mean total P(r2 = 0.95), and with mean total N(r2 = 0.91). In contrast, integral rates of photosynthesis are linked less tightly to nutrient concentration because of their simultaneous dependence on transparency. The Dillon-Rigler phosphorus loading model is extended to predict volumetric rates of photosynthesis (v and Āopt) in lakes where the N:P balance indicates control by phosphorus (N:P ⩾ 13).

https://aslopubs.onlinelibrary.wiley.com/doi/pdf/10.4319/lo.1979.24.6.1051

Nutrient dependence of primary productivity in lakes1

The validity of a recent nutrient-based modal of algal photosynthesis was tested using phosphorus-limited semi-continuous cultures of the green alga, Scenedesmus quadricauda. The data suggest that light-saturate photosynthesis (P opt, mg C · mg chla−1· h−1) is a hyperbolic function of cellular phosphorus quota (Qp), whether Qp is expressed per unit cellular chlorophyll a, per unit dry weight, or per cell number. Furthermore, the light availability during growth appears to have little effect on these hyperbolic relationships over the range of light intensities studied here (65–110 μE · m−2· s−1). These data, and data presented from other published studies, suggest that the cell quota model is a valid empirical description of the photosynthetic behavior of nutrient-limited cells.

Val H. Smith

Papers

The nitrogen and phosphorus dependence of algal biomass in lakes: An empirical and theoretical analysis1

Nutrient dependence of primary productivity in lakes1

Light and nutrient dependence of photosynthesis by algae1