Does the Redfield ratio infer nutrient limitation in the macroalga Spirogyra fluviatilis?

1. The cellular nutrient contents of microalgae, when growing at or approaching maximum rates, approximate the Redfield C : N : P (molar) ratio of 106 : 16 : 1. Deviations from this optimal ratio can be used to infer nutrient limitation of microalgal growth. However, this ratio may not be applicable to macroalgae, which are distinguished from microalgae by forming a thallus that is a discrete structure visible to the naked eye. The utility of the Redfield ratio to infer nutrient limitation of the growth of macroalgae was tested for Spirogyra fluviatilis in a field experiment conducted in tropical Australia. 2. The optimal cellular C : N : P ratio for S. fluvialitis was estimated by means of in situ nutrient addition. This was compared with S. fluvialitis cellular ratios determined from eight sites with a wide range of soluble N concentrations (<1–90 μg L^?1), a smaller range of soluble P concentrations (5–12 μg L^?1), and soluble molar N : P ratios of 0.11– 27. 3. Spirogyra fluviatilis had an optimal molar C : N : P ratio of 1800 : 87 : 1 which differs substantially from the Redfield ratio, and suggests that the latter ratio is not applicable to this macroalga. Concentrations of N and P in the river deviated from the optimal N : P ratio of 87 : 1, inferring nutrient limitation of growth. 4. C : P and C : N ratios of S. fluviatilis varied in accordance with general stoichiometric relationships for autotrophs under nutrient limitation of growth. Ratios of C : P and C : N increased, respectively, with increased severity of P‐ and N‐limitation. Additionally, C : P ratios increased with increased N : P ratios, whilst the C : N ratio increased with decreased N : P ratios. The C : N molar ratio however was an insensitive indicator of nutrient depletion compared with the C : P ratio. Under N‐limitation of growth, luxury amounts of P were stored by S. fluviatilis. 5. In aquatic environments where macroalgae are sufficiently abundant to be sampled, their cellular carbon, nitrogen and phosphorus stoichiometry can be used to infer nutrient limitation of growth when their optimal C : N : P ratio is known.     

STEADY-STATE LUXURY CONSUMPTION AND THE CONCEPT OF OPTIMUM NUTRIENT RATIOS: A STUDY WITH PHOSPHATE AND NITRATE LIMITED SELENASTRUM MINUTUM (CHLOROPHYTA)1

Selenastrum minutum (Naeg.) Collins was grown over a wide range of growth rates under phosphate or nitrate limitation with non-limiting nutrients added to great excess. This resulted in saturated luxury consumption. The relationships between growth rate and cell quota for the limiting nutrients were well described by the Droop relationship. The observed variability in N cell quota under N limitation as reflected in k_Q·Q_max^?1*, was similar in magnitude to previously reported values but k_Q·Q_max^?1* for P under P limitation was greater than previously reported for other species. These results were evaluated in light of the optimum ratio hypothesis. Our findings support previous work suggesting that the use of a single optimum ratio (k_Qi·K_Qj^?1) is inappropriate for dealing with a species growing under steady-state nutrient limitation. Under these conditions the optimum ratio should be viewed as a growth rate dependent variable. Two approaches for testing the growth rate dependency of optimum ratios are proposed. The capacity for luxury consumption differed between nutrients and was growth rate dependent. At low growth rates, the coefficient of luxury consumption (R_sat) for P was ca. four times that for N. The set of all possible relationships between N and P cell quota under these conditions was reported and these values were then used to establish the cellular N:P niche boundaries for S. minutum. Cell quotas of non-limiting nutrients were not described by the Droop equation. Analysis showed that as the cellular N:P ratio deviates from the optimum ratio, the ability of the Droop equation to describe the relationship between growth rate and non-limiting cell quotas decreases. When non-limiting nutrient cell quotas are saturated, the Droop equation appears to be invalid. Previously reported patterns of non-limiting nutrient utilization are summarized in support of this conclusion. The physiological and ecological consequences of luxury consumption and growth rate dependent optimum ratios are considered.     

Physiological integration impacts nutrient use and stoichiometry in three clonal plants under heterogeneous habitats

Physiological integration facilitates clonal plants to deal with heterogeneous resources. However, little is known about how nutrient patchiness affects its use and stoichiometry in clonal plants. We conducted an experiment with Cynodon dactylon, Glechoma longituba, and Potentilla reptans to address the effects of physiological integration on nutrient use efficiency and N:P ratios. For C. dactylon, the effects of nutrient patchiness on N use efficiency (NUE), P use efficiency (PUE), and N:P ratio were stronger in daughter ramets than in parent ramets; for G. longituba, nutrient patchiness affected PUE and N:P ratio of parent and daughter ramets, but not NUE; for P. reptans, nutrient patchiness decreased NUE, PUE, and N:P ratio, regardless of parent or daughter ramets. PUE was associated with N:P ratios in three clonal plants and this association of NUE with N:P ratios varied with species. Our findings suggest that physiological integration alters nutrient use efficiency and N:P ratios of clonal plants under patchy nutrients and that these effects are linked to clonal species identity.     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Effects of light intensity and quality on the relative N and P requirement (the optimum N:P ratio) of marine planktonic algae

The relative requirement of N and P (the optimum N:P ratio)by Dunaliella tertiolecta, Phaeodactylum tricornutum, Prymnesiumparvum and Thalassiosira pseudonana was studied under variouslight intensities and spectra. The ratio was determined as theratio of the minimum cell N and P concentrations (q₀N and q₀pwhen either nutrient was limiting. The ratio varied widely amongspecies; under light-saturation for growth (116 µEin ^m–2s^–1 it ranged from 11.8 in ^D. tertiolecta to 36.6 in P.tricornutum. The ratio appeared to be higher at a sub-saturatingintensity (24 µEin m^–2 s^–1 in all except P.tricornutum, mainly because of higher q_oN with little changein q_oP. In T. pseudonana Q_oP also increased, resulting in aninsignificant change in the ratio. The ratio varied little withinthe range of saturation intensity. Light quality affected q_oNand q_oP as well as the ratio, and the pattern of change variedfrom species to species. The optimum ratio of individual specieswas linearly correlated to their q_oN except in P. tricornutum.q_oN for all species showed a linear correlation with cell proteinconcentrations irrespective of light conditions. The changeof optimum N:P ratios in the three species thus appears to berelated to changes in cell protein contents. The ratio of carbohydratesto protein remained constant regardless of light intensity orquality and was higher in P-limited cultures. We conclude thatchanges in light regime can strongly influence algal nutrientrequirements and species interrelationships by altering theoptimum cellular N:P ratio.     

Seaweed biogeochemistry: Global assessment of C:N and C:P ratios and implications for ocean afforestation

Algal carbon-to-nitrogen (C:N) and carbon-to-phosphorus (C:P) ratios are fundamental for understanding many oceanic biogeochemical processes, such as nutrient flux and climate regulation. We synthesized literature data (444 species, >400 locations) and collected original samples from Tasmania, Australia (51 species, 10 locations) to update the global ratios of seaweed carbon-to-nitrogen (C:N) and carbon-to-phosphorus (C:P). The updated global mean molar ratio for seaweed C:N is 20 (ranging from 6 to 123) and for C:P is 801 (ranging from 76 to 4102). The C:N and C:P ratios were significantly influenced by seawater inorganic nutrient concentrations and seasonality. Additionally, C:N ratios varied by phyla. Brown seaweeds (Ochrophyta, Phaeophyceae) had the highest mean C:N of 27.5 (range: 7.6–122.5), followed by green seaweeds (Chlorophyta) of 17.8 (6.2–54.3) and red seaweeds (Rhodophyta) of 14.8 (5.6–77.6). We used the updated C:N and C:P values to compare seaweed tissue stoichiometry with the most recently reported values for plankton community stoichiometry. Our results show that seaweeds have on average 2.8 and 4.0 times higher C:N and C:P than phytoplankton, indicating seaweeds can assimilate more carbon in their biomass for a given amount of nutrient resource. The stoichiometric comparison presented herein is central to the discourse on ocean afforestation (the deliberate replacement of phytoplankton with seaweeds to enhance the ocean biological carbon sink) by contributing to the understanding of the impact of nutrient reallocation from phytoplankton to seaweeds under large-scale seaweed cultivation.     

RESPONSE OF A LENTIC PERIPHYTON COMMUNITY TO NUTRIENT ENRICHMENT AT LOW N:P RATIOS1

We examined the effect of nitrogen:phosphorus (N:P) ratios and nutrient concentrations on periphyton when nutrients (N and P) are provided in excess. A gradient of seven N:P ratios ranging from 7.5:1 to 1:7.5 and each at three absolute concentrations, was established using nutrient‐releasing substrata placed in a meso‐oligotrophic lake. Differences in total algal biovolume among nutrient ratios were significant (analysis of covariance [ANCOVA]) when P concentration was entered as the co‐variate. In addition, total algal biovolume was significantly correlated with N concentration but not P. To further evaluate the relationship between nutrient ratios and biovolume, we analyzed (using four 1‐way analysis of variances [ANOVAs]) four subsets of data defined as a series of treatments where one nutrient concentration remained relatively constant as the other changed creating different N:P ratios. Ratios of data subsets ranged from 1:1 to 7.5:1 and 1:1 to 1:7.5 with low and high concentrations of both series. Only diatom biovolume varied with ratio but these differences are most likely related to increased green algal abundance. Species richness and diversity differed among N:P ratios (ANCOVA) when P concentration was used as the co‐variate. Stigeoclonium tenue (Ag.) Gomont, which generally accounted for the increase in green algal abundance, varied with nutrient ratio (ANCOVA) when P was the co‐variate. Based on the ANCOVAs, correlations, and one‐way ANOVAs, periphyton in this system appears to be affected by N concentration but not by N:P treatment ratios under nutrient‐rich conditions. When compared with previous studies, these data also suggest that the response of periphyton to in situ treatments constructed with nutrient‐releasing substrata vary between years.     

Variation in nitrogen and phosphorus concentrations of wetland plants

The use of nutrient concentrations in plant biomass as easily measured indicators of nutrient availability and limitation has been the subject of a controversial debate. In particular, it has been questioned whether nutrient concentrations are mainly species' traits or mainly determined by nutrient availability, and whether plant species have similar or different relative nutrient requirements. This review examines how nitrogen and phosphorus concentration and the N:P ratio in wetland plants vary among species and sites, and how they are related to nutrient availability and limitation. We analyse data from field studies in European non-forested wetlands, from fertilisation experiments in these communities and from growth experiments with wetland plants. Overall, the P concentration was more variable than the N concentration, while variation in N:P ratios was intermediate. Field data showed that the N concentration varies more among species than among sites, whereas the N:P ratio varies more among sites than among species, and the P concentration varies similarly among both. Similar patterns of variation were found in fertilisation experiments and in growth experiments under controlled nutrient supply. Nutrient concentrations and N:P ratios in the vegetation were poorly correlated with various measures of nutrient availability in soil, but they clearly responded to fertilisation in the field and to nutrient supply in growth experiments. In these experiments, biomass N:P ratios ranged from 3 to 40 and primarily reflected the relative availabilities of N and P, although N:P ratios of plants grown at the same nutrient supply could vary three-fold among species. The effects of fertilisation with N or P on the biomass production of wetland vegetation were well related to the N:P ratios of the vegetation in unfertilised plots, but not to N or P concentrations, which supports the idea that N:P ratios, rather than N or P concentrations, indicate the type of nutrient limitation. However, other limiting or stressing factors may influence N:P ratios, and the responses of individual plant species to fertilisation cannot be predicted from their N:P ratios. Therefore, N:P ratios should only be used to assess which nutrient limits the biomass production at the vegetation level and only when factors other than N or P are unlikely to be limiting.     

Ontogeny, diet shifts, and nutrient stoichiometry in fish

Most stoichiometric models do not consider the importance of ontogenetic changes in body nutrient composition and excretion rates. We quantified ontogenetic variation in stoichiometry and diet in gizzard shad, Dorosoma cepedianum , an omnivorous fish with a pronounced ontogenetic diet shift; and zebrafish, Danio rerio, grown in the lab with a constant diet. In both species, body stoichiometry varied considerably along the life cycle. Larval gizzard shad and zebrafish had higher molar C:P and N:P ratios than larger fish. Variation in body nutrient ratios was driven mainly by body P, which increased with size. Gizzard shad body calcium content was highly correlated with P content, indicating that ontogenetic P variation is associated with bone formation. Similar trends in body stoichiometry of zebrafish, grown under constant diet in the laboratory, suggest that ontogeny (e.g. bone formation) and not diet shift is the main factor affecting fish body stoichiometry in larval and juvenile stages. The N:P ratio of nutrient excretion also varied ontogenetically in gizzard shad, but the decline from larvae to juveniles appears to be largely associated with variation in the N:P of alternative food resources (zooplankton vs detritus) rather than by fish body N:P. Furthermore, the N:P ratio of larval gizzard shad excretion appears to be driven more by the N:P ratio at which individuals allocate nutrients to growth, more so than static body N:P, further illustrating the need to consider ontogenetic variation. Our results thus show that fish exhibit considerable ontogenetic variation in body stoichiometry, driven by an inherent increase in the relative allocation of P to bones, whereas ontogenetic variation in excretion N:P ratio of gizzard shad is driven more by variation in food N:P than by body N:P.     

The C:N:P stoichiometry of organisms and ecosystems in a changing world: A review and perspectives

This study examined the literature in ISI Web of Science to identify the effects that the main drivers of global change have on the nutrient concentrations and C:N:P stoichiometry of organisms and ecosystems, and examined their relationship to changes in ecosystem structure and function. We have conducted a meta-analysis by comparing C:N:P ratios of plants and soils subjected to elevated [CO₂] with those subjected to ambient [CO₂]. A second meta-analysis compared the C:N:P ratios of plants and soils that received supplemental N to simulate N deposition and those that did not receive supplemental N. On average, an experimental increase in atmospheric [CO₂] increased the foliar C:N ratios of C3 grasses, forbs, and woody plants by 22%, but the foliar ratios of C4 grasses were unaffected. This trend may be enhanced in semi-arid areas by the increase in droughts that have been projected for the coming decades which can increase leaf C:N ratios. The available studies show an average 38% increase in foliar C:P ratios in C3 plants in response to elevated atmospheric [CO₂], but no significant effects were observed in C4 grasses. Furthermore, studies that examine the effects of elevated atmospheric [CO₂] on N:P ratio (on a mass basis) are warranted since its response remains elusive. N deposition increases the N:P ratio in the plants of terrestrial and freshwater ecosystems, and decreases plants and organic soil C:N ratio (25% on average for C3 plants), reducing soil and water N₂ fixation capacity and ecosystem species diversity. In contrast, in croplands subjected to intense fertilization, mostly, animal slurries, a reduction in soil N:P ratio can occur because of the greater solubility and loss of N. In the open ocean, there are experimental observations showing an ongoing increase in P-limited areas in response to several of the factors that promote global change, including the increase in atmospheric [CO₂] which increases the demand for P, the warming effect that leads to an increase in water column stratification, and increases in the N:P ratio of atmospheric inputs. Depending on the type of plant and the climate where it grows, warming can increase, reduce, or have no effect on foliar C:N ratios. The results suggest that warming and drought can increase C:N and C:P ratios in warm-dry and temperate-dry terrestrial ecosystems, especially, when high temperatures and drought coincide. Advances in this topic are a challenge because changes in stoichiometric ratios can favour different types of species and change ecosystem composition and structure.     

Stoichiometry of nutrient recycling by vertebrates in a tropical stream: linking species identity and ecosystem processes

Ecological stoichiometry offers a framework for predicting how animal species vary in recycling nutrients, thus providing a mechanism for how animal species identity mediates ecosystem processes. Here we show that variation in the rates and ratios at which 28 vertebrate species (fish, amphibians) recycled nitrogen (N) and phosphorus (P) in a tropical stream supports stoichiometry theory. Mass-specific P excretion rate varied 10-fold among taxa and was negatively related to animal body P content. In addition, the N : P ratio excreted was negatively related to body N : P. Body mass (negatively related to excretion rates) explained additional variance in these excretion parameters. Body P content and P excretion varied much more among taxonomic families than among species within families, suggesting that familial composition may strongly influence ecosystem-wide nutrient cycling. Interspecific variation in nutrient recycling, mediated by phylogenetic constraints on stoichiometry and allometry, illustrates a strong linkage between species identity and ecosystem function.     

Responses of wetland graminoids to the relative supply of nitrogen and phosphorus

The biomass production of wetland vegetation can be limited by nitrogen or phosphorus. Some species are most abundant in N-limited vegetation, and others in P-limited vegetation, possibly because growth-related traits of these species respond differently to N versus P supply. Two growth experiments were carried out to examine how various morphological and physiological traits respond to the relative supply of N and P, and whether species from sites with contrasting nutrient availability respond differently. In experiment 1, four Carex species were grown in nutrient solutions at five N:P supply ratios (1.7, 5, 15, 45, 135) combined with two levels of supply (geometric means of N and P supply). In experiment 2, two Carex and two grass species were grown in sand at the same .ve N:P supply ratios combined with three levels of supply and two light intensities (45% or 5% daylight). After 12-13 weeks of growth, plant biomass, allocation, leaf area, tissue nutrient concentrations and rates and nutrient uptake depended signi.cantly on the N:P supply ratio, but the type and strength of the responses differed among these traits. The P concentration and the N:P ratio of shoots and roots as well as the rates of N and P uptake were mainly determined by the N:P supply ratio; they showed little or no dependence on the supply level and relatively small interspeci.c variation. By contrast, the N concentration, root mass ratio, leaf dry matter content and speci.c leaf area were only weakly related to the N:P supply ratio; they mainly depended on plant species and light, and partly on overall nutrient supply. Plant biomass was determined by all factors together. Within a level of light and nutrient supply, biomass was generally maximal (i.e. co-limited by N and P) at a N:P supply ratio of 15 or 45. All species responded in a similar way to the N:P supply ratio. In particular, the grass species Phalaris arundinacea and Molinia caerulea showed no differences in response that could clearly explain why P. arundinacea tends to invade P-rich (N-limited) sites, and M. caerulea P-limited sites. This may be due to the short duration of the experiments, which investigated growth and nutrient acquisition but not nutrient con­servation.     

GROWTH RATE DEPENDENT OPTIMUM RATIOS IN SELENASTRUM MINUTUM (CHLOROPHYTA): IMPLICATIONS FOR COMPETITION,COEXISTENCE AND STABILITY IN PHYTOPLANKTON COMMUNITIES 12

Steady-State growth equations predict that the optimum C:P ratio (R) of Selenastrum minutum (Naeq.) Collins should change by a factor of 20 over the growth range of this organism. Chemostat cultures were established at fixed inflow C:P ratios chosen such that a transition from carbon to phosphorus limitation should occur solely as a result of increasing the steady-state growth rate. Measurements of residual dissolved inorganic carbon (DIC), cellular C:P, the kinetics of photosynthesis with respect to [DIC] and the response of culture biomass lo DIC or K₂HPO₄ additions were obtained. These results show that optimum ratios are growth rate dependent and that this dependency can be predicted based on steady-stale algal growth equations. A theoretical analysis was undertaken evaluating the range of growth rate dependent changes in the optimum ratio which could be expected for different nutrient pairs. Further analysis showed that, under certain conditions, the growth rate dependence of the optimum ratio may alter the breadth of zones of stable coexistence between species and allow for either the formation or complete elimination of such zones.     

Diel variation of the cellular carbon to nitrogen ratio of Chlorella autotrophica (Chlorophyta) growing in phosphorus‐ and nitrogen‐limited continuous cultures

We investigated the relationship between daily growth rates and diel variation of carbon (C) metabolism and C to nitrogen (N) ratio under P‐ and N‐limitation in the green algae Chlorella autotrophica. To do this, continuous cultures of C. autotrophica were maintained in a cyclostat culture system under 14:10 light:dark cycle over a series of P‐ and N‐limited growth rates. Cell abundance, together with cell size, as reflected by side scatter signal from flow cytometric analysis demonstrated a synchronized diel pattern with cell division occurring at night. Under either type of nutrient limitation, the cellular C:N ratio increased through the light period and decreased through the dark period over all growth rates, indicating a higher diel variation of C metabolism than that of N. Daily average cellular C:N ratios were higher at lower dilution rates under both types of nutrient limitation but cell enlargement was only observed at lower dilution rates under P‐limitation. Carbon specific growth rates during the dark period positively correlated with cellular daily growth rates (dilution rates), with net loss of C during night at the lowest growth rates under N‐limitation. Under P‐limitation, dark C specific growth rates were close to zero at low dilution rates but also exhibited an increasing trend at high dilution rates. In general, diel variations of cellular C:N were low when dark C specific growth rates were high. This result indicated that the fast growing cells performed dark C assimilation at high rates, hence diminished the uncoupling of C and N metabolism at night.     

Ecoenzymatic stoichiometry of stream sediments with comparison to terrestrial soils

The kinetics and elemental composition of cellular units that mediate production and respiration are the basis for the metabolic and stoichiometric theories of ecological organization. This theoretical framework extends to the activities of microbial enzymes released into the environment (ecoenzymes) that mediate the release of assimilable substrate from detrital organic matter. In this paper, we analyze the stoichiometry of ecoenzymatic activities in the surface sediments of lotic ecosystems and compare those results to the stoichiometry observed in terrestrial soils. We relate these ecoenzymatic ratios to energy and nutrient availability in the environment as well as microbial elemental content and growth efficiency. The data, collected by US Environmental Protection Agency, include the potential activities of 11 enzymes for 2,200 samples collected across the US, along with analyses of sediment C, N and P content. On average, ecoenzymatic activities in stream sediments are 2–5 times greater per gC than those of terrestrial soils. Ecoenzymatic ratios of C, N and P acquisition activities support elemental analyses showing that microbial metabolism is more likely to be C-limited than N or P-limited compared to terrestrial soils. Ratios of hydrolytic to oxidative activities indicate that sediment organic matter is more labile than soil organic matter and N acquisition is less dependent on humic oxidation. The mean activity ratios of glycosidases and aminopeptidases reflect the environmental abundance of their respective substrates. For both freshwater sediments and terrestrial soils, the mean C:nutrient ratio of microbial biomass normalized to growth efficiency approximates the mean ecoenzymatic C:nutrient activity ratios normalized to environmental C:nutrient abundance. This relationship defines a condition for biogeochemical equilibrium consistent with stoichiometric and metabolic theory.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

The relationship between relative growth rate and whole-plant C:N:P stoichiometry in plant seedlings grown under nutrient-enriched conditions

Aims Recent theories indicate that N is more in demand for plant growth than P; therefore, N concentration and N : C and N : P ratios are predicted to be positively correlated with relative growth rate (RGR) in plants under nutrient-enriched conditions. This prediction was tested in this study.Methods We examined the whole-plant concentrations of C, N and P and RGR, as well as the relationship between RGR and the concentrations and the ratios of N : C, P : C and N : P, for different harvest stages (the days after seed germination) of the seedlings of seven shrub species and four herbaceous species grown in N and P non-limiting conditions. The relationships among plant size, nutrient concentrations and ratios were subsequently determined.Important findings RGR was positively correlated with N concentration and the ratios of N : P and N : C when the data were pooled for all species and for each shrub species, but not for individual herbaceous species. However, the relationship between RGR and P concentration and P : C was not significantly correlated for either shrubs or herbs. The variation of N among harvest stages and species was much greater than that of P, and the variation in N : P ratio was determined primarily by changes in N concentration. The shrub species differed from the herbaceous species in their N and P concentrations, nutrient ratios and in intraspecific relationships between RGR and nutrient ratios. These differences possibly reflect differences in the capacity for P storage and biomass allocation patterns. In general, our data support recent theoretical predictions regarding the relationship between RGR and C : N : P stoichiometry, but they also show that species with different life forms differ in the relationships among RGR and C : N : P stoichimetries.     

Allometric and trait‐based patterns in parasite stoichiometry

We measured the elemental content (%C, N and P) and ratios (C:N, C:P, N:P) of a diverse assemblage of parasitic helminths to ask whether taxonomy or traits were related to stoichiometric variation among species. We sampled 27 macroparasite taxa, spanning four phyla, infecting vertebrate and invertebrate hosts from freshwater ecosystems in New Jersey. Macroparasites varied widely in elemental content, exhibiting 4.7‐fold variation in %N, 4.6‐fold variation in %P, and 11.5‐fold variation in N:P. Across all species, parasite %P scaled negatively and C:P scaled positively with body size. Similar relationships between parasite P content and body size occurred at the phylum level and within individual species. The allometric scaling of P across species supports the growth rate hypothesis, which predicts that smaller taxa require more P to support relatively higher growth rates. Life cycle stage was related to %N and C:N, with non‐reproductive parasite stages lower in %N and higher in C:N than actively reproducing parasites. Parasite phylum, functional feeding group, and trophic level did not explain elemental variation among species. Organismal stoichiometry is linked to ecological function, and wide variation in macroparasite stoichiometry likely generates diverse patterns in host–parasite nutrient dynamics and variable relationships between parasitism and nutrient cycling.     

天童常绿阔叶林演替系列植物群落的N∶P化学计量特征

 土壤氮磷养分对植物生长的限制性可通过植被的N∶P化学计量特征来反映。该研究以常绿阔叶林演替系列为对象,将N∶P作为诊断指标,揭示 常绿阔叶林次生演替过程中植物群落的N∶P化学计量特征和养分限制作用。结果显示：1)物种水平的N∶P大小不一,但演替系列总体的变化特 征表现出了较高的一致性。2）在群落水平上,次生演替初期的灌草丛N∶P极小(7.38),远远低于14,当演替进入灌丛阶段,N∶P 显著增高到 19.96,在进入演替中期的针叶林(14.29)和针阔混交林(14.21)时,N∶P显著下降到 14～16之间,演替中后期的木荷(Schima superba)群落 (18.77)和栲树(Castanopsis fargesii)群落(20.13)的N∶P发生了显著的升高过程 。根据以往对N∶P临界值的确定,可以认为,常绿阔叶林次 生演替初期的植物群落生产力主要受到氮素的限制作用;演替中期的针叶林和针阔混交林主要受氮磷的共同限制,但以氮素的限制作用更为强 烈;演替中后期植物群落主要受到土壤磷素的限制作用。     

Carbon,Nitrogen, Phosphorus,and Potassium Stoichiometry in an Ombrotrophic Peatland Reflects Plant Functional Type

Ombrotrophic bog peatlands are nutrient-deficient systems and important carbon (C) sinks yet the stoichiometry of nitrogen (N), phosphorus (P) and potassium (K), essential for plant growth and decomposition, has rarely been studied. We investigated the seasonal variation in C, N, P, and K concentrations and their stoichiometric ratios in photosynthetically active tissues of 14 species belonging to five plant functional types (PFTs) (mosses, deciduous trees/shrubs, evergreen shrubs, graminoids, and forb) at Mer Bleue bog, an ombrotrophic peatland in eastern Ontario, Canada. Although we observed variations in stoichiometry among PFTs at peak growing season, there was convergence of C:N:P:K to an average mass ratio of 445:14:1:9, indicating N and P co-limitation. Nitrogen, P, and K concentrations and stoichiometric ratios showed little seasonal variation in mosses, evergreens, and graminoids, but in forb and deciduous species were the largest in spring and decreased throughout the growing season. Variations in nutrient concentrations and stoichiometric ratios among PFTs were greater than seasonal variation within PFTs. Plants exhibit N and P co-limitation and adapt to extremely low nutrient availability by maintaining small nutrient concentrations in photosynthetically active tissues, especially for evergreen shrubs and Sphagnum mosses. Despite strong seasonal variations in nutrient availabilities, few species show strong seasonal variation in nutrient concentrations, suggesting a strong stoichiometric homeostasis at Mer Bleue bog.