Responses of leaf C:N:P stoichiometry to water supply in the desert shrub Zygophyllum xanthoxylum

• Based on the elemental composition of major biochemical molecules associated with different biological functions, the ‘growth rate hypothesis’ proposed that organisms with a higher growth rate would be coupled to lower C:N, especially lower C:P and N:P ratios. However, the applicability of the growth rate hypothesis for plants is unclear, especially for shrubs growing under different water supply.
• We performed an experiment with eight soil moisture levels (soil water content: 4%, 6%, 8%, 13%, 18%, 23%, 26% and 28%) to evaluate the effects of water availability on leaf C:N:P stoichiometry in the shrub Zygophyllum xanthoxylum.
• We found that leaves grew slowly and favored accumulation of P over C and N under both high and low water supply. Thus, leaf C:P and N:P ratios were unimodally related to soil water content, in parallel with individual leaf area and mass. As a result, there were significant positive correlations between leaf C:P and N:P with leaf growth (u).
• Our result that slower‐growing leaves had lower C:P and N:P ratios does not support the growth rate hypothesis, which predicted a negative association of N:P ratio with growth rate, but it is consistent with recent theoretical derivations of growth–stoichiometry relations in plants, where N:P ratio is predicted to increase with increasing growth for very low growth rates, suggesting leaf growth limitation by C and N rather than P for drought and water saturation.

     

N : P stoichiometry and protein : RNA ratios in vascular plants: an evaluation of the growth-rate hypothesis

The growth-rate hypothesis states that fast-growing organisms need relatively more phosphorus-rich RNA to support rapid rates of protein synthesis, and therefore predicts, within and among taxa, increases in RNA and phosphorus content (relative to protein and nitrogen content) with increased growth rate. Here, we present a test of this hypothesis in vascular plants. We determined nitrogen : phosphorus ratios and protein : RNA ratios in pines growing at different rates due to nutrient conditions. In general, when comparing leaves of the same species at low and high growth rates, the faster-growing plants had higher RNA content, higher %N and %P, and lower protein : RNA ratios, but not consistently lower N : P ratios. We found no link between growth rate and foliar N : P or protein : RNA when comparing multiple species of different inherent growth rates. We conclude that plants adjust the balance of protein and RNA to favour either speed or efficiency of protein synthesis, but this balance does not alone dictate leaf stoichiometry.     

DIAZOTROPHIC GROWTH OF THE MARINE CYANOBACTERIUM TRICHODESMIUM IMS101 IN CONTINUOUS CULTURE: EFFECTS OF GROWTH RATE ON N2‐FIXATION RATE,BIOMASS, AND C:N:P STOICHIOMETRY1

Trichodesmium N₂ fixation has been studied for decades in situ and, recently, in controlled laboratory conditions; yet N₂‐fixation rate estimates still vary widely. This variance has made it difficult to accurately estimate the input of new nitrogen (N) by Trichodesmium to the oligotrophic gyres of the world ocean. Field and culture studies demonstrate that trace metal limitation, phosphate availability, the preferential uptake of combined N, light intensity, and temperature may all affect N₂ fixation, but the interactions between growth rate and N₂ fixation have not been well characterized in this marine diazotroph. To determine the effects of growth rate on N₂ fixation, we established phosphorus (P)–limited continuous cultures of Trichodesmium, which we maintained at nine steady‐state growth rates ranging from 0.27 to 0.67 d^?1. As growth rate increased, biomass (measured as particulate N) decreased, and N₂‐fixation rate increased linearly. The carbon to nitrogen ratio (C:N) varied from 5.5 to 6.2, with a mean of 5.8 ± 0.2 (mean ± SD, N = 9), and decreased significantly with growth rate. The N:P ratio varied from 23.4 to 45.9, with a mean of 30.5 ± 6.6 (mean ± SD, N = 9), and remained relatively constant over the range of growth rates studied. Relative constancy of C:N:P ratios suggests a tight coupling between the uptake of these three macronutrients and steady‐state growth across the range of growth rates. Our work demonstrates that growth rate must be considered when planning studies of the effects of environmental factors on N₂ fixation and when modeling the impact of Trichodesmium as a source of new N to oligotrophic regions of the ocean.     

刨花楠不同相对生长速率下林木叶片碳氮磷的适应特征

分别对9年生与13年生刨花楠林木叶片氮磷养分之间关系及林木生物量相对生长速率与叶片碳氮磷化学计量比关系进行分析,探讨不同相对生长速率下的林木叶片N、P养分适应特征,并检验相对生长速率假说理论对刨花楠树种的适应性。结果表明:两种年龄刨花楠林木生物量相对生长速率、叶片C、N、P含量及其计量比值均存在显著差异;同一年龄的林木叶片N、P之间存在显著相关性,二者具有协同相关性;9年生林木叶片P含量及C∶P、N∶P与生物量相对生长速率呈二次曲线相关,而13年生林木叶片N、P含量及C∶N、C∶P、N∶P则与生物量相对生长速率均呈线性相关。研究表明,在能满足植物生长所需养分供给的土壤环境中,叶片N、P含量与林木相对生长速率间呈线性正相关,但当土壤中养分供应满足不了植物高速生长时,植物则会对有限的养分资源进行适应性调整。     

C : N : P stoichiometry of dominant riparian trees and arthropods along the Middle Rio Grande

1. We examined the role of flooding on the leaf nutrient content of riparian trees by comparing the carbon : nitrogen : phosphorus (C : N : P) ratio of leaves and litter of Rio Grande cottonwood (Populus deltoides ssp. wislizenii) in flood and non‐flood sites along the Middle Rio Grande, NM, U.S.A. The leaf C : N : P ratio was also examined for two non‐native trees, saltcedar (Tamarix chinensis) and Russian olive (Elaeagnus angustifolia), and six species of dominant riparian arthropods. 2. Living leaves and leaf litter of cottonwoods at flood sites had a significantly lower leaf N : P ratio and higher %P compared with leaves and litter at non‐flood sites. A non‐flood site downstream from wastewater effluent had a significantly lower litter C : N ratio than all other sites, suggesting N fertilisation through ground water. The non‐native trees, saltcedar and Russian olive, had higher mean leaf N content, N : P ratio, and lower C : N ratio compared with cottonwoods across study sites. 3. Riparian arthropods ranged from 5.2 to 7.1 for C : N ratio, 56–216 for C : P ratio, and 8.9–34 for N : P ratio. C content ranged from 25 to 52% of dry mass, N content from 4.7 to 10.8%, and P content from 0.59 to 1.2%. Differences in stoichiometry between high C : nutrient leaf litter and low C : nutrient invertebrates suggests possible food‐quality constraints for detritivores. 4. These results suggest that spatial and temporal variation in the C : N : P ratio of cottonwood leaves and leaf litter is influenced by surface and subsurface hydrologic connection within the floodplain. Reach‐scale variation in the elemental composition of riparian organic matter inputs may have important implications for decomposition, nutrient cycling, and food webs in river floodplain systems.     

Elemental and macromolecular plasticity of Chlamydomonas reinhardtii (Chlorophyta) in response to resource limitation and growth rate

With the ongoing differential disruption of the biogeochemical cycles of major elements that are essential for all life (carbon, nitrogen, and phosphorus), organisms are increasingly faced with a heterogenous supply of these elements in nature. Given that photosynthetic primary producers form the base of aquatic food webs, impacts of changed elemental supply on these organisms are particularly important. One way that phytoplankton cope with the differential availability of nutrients is through physiological changes, resulting in plasticity in macromolecular and elemental biomass composition. Here, we assessed how the green alga Chlamydomonas reinhardtii adjusts its macromolecular (e.g., carbohydrates, lipids, and proteins) and elemental (C, N, and P) biomass pools in response to changes in growth rate and the modification of resources (nutrients and light). We observed that Chlamydomonas exhibits considerable plasticity in elemental composition (e.g., molar ratios ranging from 124 to 971 for C:P, 4.5 to 25.9 for C:N, and 15.1 to 61.2 for N:P) under all tested conditions, pointing to the adaptive potential of Chlamydomonas in a changing environment. Exposure to low light modified the elemental and macromolecular composition of cells differently than limitation by nutrients. These observed differences, with potential consequences for higher trophic levels, included smaller cells, shifts in C:N and C:P ratios (due to proportionally greater N and P contents), and differential allocation of C among macromolecular pools (proportionally more lipids than carbohydrates) with different energetic value. However, substantial pools of N and P remained unaccounted for, especially at fast growth, indicating accumulation of N and P in forms we did not measure.     

STOICHIOMETRIC RESPONSES OF PHYTOPLANKTON SPECIES TO THE INTERACTIVE EFFECT OF NUTRIENT SUPPLY RATIOS AND GROWTH RATES1

Three species of phytoplankton, Rhodomonas sp., Phaeodactylum tricornutum Bohlin, and Isochrysis galbana Parke, were cultivated in semicontinuous culture to analyze the response of carbon (C):nitrogen (N):phosphorus (P) stoichiometry to the interactive effect of five N:P supply ratios and four growth rates (dilution rates). The relationship between cellular N and P quotas and growth rates fits well to both the Droop and Ågren’s functions for all species. We observed excess uptake of both N and P in the three species. N:P biomass ratios showed a significant positive relationship with N:P supply ratios across the entire range of growth rates, and N:P biomass ratios converged to an intermediate value independent of N:P supply ratios at higher growth rates. The effect of growth rates on N:P biomass ratios was positive at lower N:P supply ratios, but negative at higher N:P supply ratios for both Rhodomonas sp. and I. galbana, while for P. tricornutum this effect was negative at all N:P supply ratios. A significant interactive effect of N:P supply ratios and growth rates on N:P biomass ratios was found in both Rhodomonas sp. and P. tricornutum, but not in I. galbana. Our results suggest that Ågren’s functions may explain the underlying biochemical principle for the Droop model. The parameters in the Droop and Ågren’s functions can be useful indications of algal succession in the phytoplankton community in changing oceans.     

Parasite and host elemental content and parasite effects on host nutrient excretion and metabolic rate

Ecological stoichiometry uses the mass balance of elements to predict energy and elemental fluxes across different levels of ecological organization. A specific prediction of ecological stoichiometry is the growth rate hypothesis (GRH), which states that organisms with faster growth or reproductive rates will require higher phosphorus content for nucleic acid and protein synthesis. Although parasites are found ubiquitously throughout ecosystems, little is understood about how they affect nutrient imbalances in ecosystems. We (1) tested the GRH by determining the carbon (C), nitrogen (N), and phosphorus (P) content of parasitic trematodes and their intermediate host, the freshwater snail Elimia livescens, and (2) used this framework to determine the trematode effects on host nutrient excretion and metabolism. Snail and parasite tissues were analyzed for elemental content using a CHN analyzer and soluble reactive phosphorus (SRP) methods. Ammonium and SRP assays were used to estimate N and P excretion rates. A respirometer was used to calculate individual snail metabolism. Trematode tissues contained lower C:P and N:P (more P per unit C and N) than the snail tissues. Snail gonadal tissues more closely resembled the elemental content of parasite tissues, although P content was 13% higher in the gonad than the trematode tissues. Despite differences in elemental content, N and P excretion rates of snails were not affected by the presence of parasites. Parasitized snails maintained faster metabolic rates than nonparasitized snails. However, the species of parasite did not affect metabolic rate. Together, this elemental imbalance between parasite and host, and the altered metabolic rate of infected snails may lead to broader parasite effects in stream ecosystems.     

Growth rate–stoichiometry couplings in diverse biota

Biological stoichiometry provides a mechanistic theory linking cellular and biochemical features of co‐evolving biota with constraints imposed by ecosystem energy and nutrient inputs. Thus, understanding variation in biomass carbon : nitrogen : phosphorus (C : N : P) stoichiometry is a major priority for integrative biology. Among various factors affecting organism stoichiometry, differences in C : P and N : P stoichiometry have been hypothesized to reflect organismal P‐content because of altered allocation to P‐rich ribosomal RNA at different growth rates (the growth rate hypothesis, GRH). We tested the GRH using data for microbes, insects, and crustaceans and we show here that growth, RNA content, and biomass P content are tightly coupled across species, during ontogeny, and under physiological P limitation. We also show, however, that this coupling is relaxed when P is not limiting for growth. The close relationship between P and RNA contents indicates that ribosomes themselves represent a biogeochemically significant repository of P in ecosystems and that allocation of P to ribosome generation is a central process in biological production in ecological systems.     

Biogeographic bases for a shift in crop C : N : P stoichiometries during domestication

We lack both a theoretical framework and solid empirical data to understand domestication impacts on plant chemistry. We hypothesised that domestication increased leaf N and P to support high plant production rates, but biogeographic and climate patterns further influenced the magnitude and direction of changes in specific aspects of chemistry and stoichiometry. To test these hypotheses, we used a data set of leaf C, N and P from 21 herbaceous crops and their wild progenitors. Domestication increased leaf N and/or P for 57% of the crops. Moreover, the latitude of the domestication sites (negatively related to temperature) modulated the domestication effects on P (+), C (?), N : P (?) and C : P (?) ratios. Further results from a litter decomposition assay showed that domestication effects on litter chemistry affected the availability of soil N and P. Our findings draw attention to evolutionary effects of domestication legacies on plant and soil stoichiometry and related ecosystem services (e.g. plant yield and soil fertility).     

Aridity Decouples C:N:P Stoichiometry Across Multiple Trophic Levels in Terrestrial Ecosystems

Increases in aridity forecasted by the end of this century will decouple the cycles of soil carbon (C), nitrogen (N) and phosphorus (P) in drylands—the largest terrestrial biome on Earth. Little is known, however, about how changes in aridity simultaneously affect the C:N:P stoichiometry of organisms across multiple trophic levels. It is imperative that we understand how aridity affects ecological stoichiometry so that we can develop strategies to mitigate any effects of changing climates. We characterized the C, N, P concentration and stoichiometry of soils, autotrophs (trees, N-fixing shrubs, grasses and mosses) and heterotrophs (microbes and ants) across a wide aridity gradient in Australia. Our results suggest that increases in aridity by the end of this century may alter the C:N:P stoichiometry of heterotrophs (ants and microbes), non-woody plants and in soil, but will not affect that one from woody plants. In particular, increases in aridity were positively related to C:P and N:P ratios in microbes and ants, negatively related to concentration of C, and the C:N and C:P ratios in mosses and/or short grasses, and not related to the C:N:P stoichiometry of either shrubs or trees. Because of the predominant role of C:N:P stoichiometry in driving nutrient cycling, our findings provide useful contextual information to determine ecological responses in a drier world.     

Growth rate of <Emphasis Type="Italic">Daphnia</Emphasis> feeding on seston in a Siberian reservoir: the role of essential fatty acid

Specific growth rates (based on biomass increment in unit time) of Daphnia fed natural reservoir seston from a eutrophic Siberian reservoir were studied during four vegetation seasons in a laboratory flow-through system. Concentrations of particulate C, N, and P in reservoir seston <115 μm were comparatively high, 1.9, 0.2, and 0.05 mg l⁻¹, respectively; maximum C:P ratio was 259 mol:mol. According to conventional thresholds, concentrations of elements and their stoichiometry did not limit the growth rate of Daphnia. Eicosapentaenoic acid (EPA) in seston significantly effected the growth rates at concentrations up to ∼13 μg l⁻¹ (regression ANOVAR F-test value was 7.91), but not above this concentration. Thus, we consider this concentration of EPA in seston <115 μm to be the limiting concentration of EPA for the growth of Daphnia (longispina group), i.e., below this concentration EPA was the best single predictor of the growth of Daphnia.     

GROWTH RATE VARIATION IN THE N:P REQUIREMENT RATIO OF PHYTOPLANKTON1

Theoretical considerations predict that the cell N:P ratio at transition from nitrogen limitation to phosphorus limitation of phytoplankton growth (critical ratio, R_c) varies, as a function of population growth rate. This prediction is confirmed by experimental, data from the literature along with new experimental data for the marine, prymnesiophyte Pavlova lutheri (Droop) Green. R_c passes through a maximum at intermediate growth rates for the three phytoplankton species for which data, are available, but there is significant interspecific variability in its value. There is no theoretical or experimental evidence to support the idea that the ratio of subsistence N and P cell quotas is equal to R_c over the range of growth rates, or that the subsistence quota ratio equals the ratio of the N and P cell quotas minus a storage fraction. Examination of N:P composition ratios can be used to determine which nutrient is limiting, but cannot be used to determine relative growth rates or competitive advantage between species limited by the same nutrient. Growth rates are determined by environmental conditions and by the cell quota of the limiting nutrient, without reference to the cell quota of the non-limiting nutrient.     

Growth and nutrient dynamics of beech (Fagus sylvatica L.) seedlings in acid soils

The relative uptake rates of N, P, K, S, Ca, Mg, Fe, Mn, Zn, Cu, and Al were estimated in beech seedlings pot cultured in the field in six acid soils (treatments). The relative uptake rates were compared with the relative growth rates. The relative uptake rates of N, K and Ca agreed well with the growth rates of the seedlings irrespective of widely differing soil conditions (acid sand-clayey till, pH 4–6). The relative uptake rates of P, Fe, and Al differed from that predicted by the growth rate. The uptake rates of Fe and Al were highest at the lowest growth rates, and the P uptake rate was lower than the growth rate in these treatments. Thus the P availability probably limited growth in an eluvial (E) horizon of a podzol, and possibly in the illuvial (B) horizon of a podzol and in an acid clayey till (Dystric Cambisol). Low P uptake was associated with a tendency towards higher relative root growth rates. In terms of the concept of steady state nutrition the high relative root growth rate in some treatments may be interpreted as an acclimation to low P supply. The P limitation seemed to be related to interactions among Fe, Al and organic compounds of the soil solution.FAX no: +4646104423     

Mixotrophy everywhere on land and in water: the grand écart hypothesis

There is increasing awareness that many terrestrial and aquatic organisms are not strictly heterotrophic or autotrophic but rather mixotrophic. Mixotrophy is an intermediate nutritional strategy, merging autotrophy and heterotrophy to acquire organic carbon and/or other elements, mainly N, P or Fe. We show that both terrestrial and aquatic mixotrophs fall into three categories, namely necrotrophic (where autotrophs prey on other organisms), biotrophic (where heterotrophs gain autotrophy by symbiosis) and absorbotrophic (where autotrophs take up environmental organic molecules). Here we discuss their physiological and ecological relevance since mixotrophy is found in virtually every ecosystem and occurs across the whole eukaryotic phylogeny, suggesting an evolutionary pressure towards mixotrophy. Ecosystem dynamics tend to separate light from non‐carbon nutrients (N and P resources): the biological pump and water stratification in aquatic ecosystems deplete non‐carbon nutrients from the photic zone, while terrestrial plant successions create a canopy layer with light but devoid of non‐carbon soil nutrients. In both aquatic and terrestrial environments organisms face a grand écart (dancer's splits, i.e., the need to reconcile two opposing needs) between optimal conditions for photosynthesis vs. gain of non‐carbon elements. We suggest that mixotrophy allows adaptation of organisms to such ubiquist environmental gradients, ultimately explaining why mixotrophic strategies are widespread.     

Linking elements to biochemicals: effects of nutrient supply ratios and growth rates on fatty acid composition of phytoplankton species

Three species of marine phytoplankton, Rhodomonas sp., Isochrysis galbana Parke, and Phaeodactylum tricornutum Bohlin, were cultivated in semicontinuous cultures to test biochemical responses (fatty acids; FAs) to five nitrogen (N):phosphorus (P) supply ratios and four growth rates (dilution rates). The characteristic FA profile was observed for each algal species (representing particular algal class), which remained relatively stable across the entire ranges of N:P supply ratios and growth rates. For all species, significant direct effects of N:P supply ratios on FAs were found at lower growth rates. The highest saturated and monounsaturated fatty acid (SFA and MUFA) contents were observed under N deficiency at the lowest growth rate in all three species, while responses of polyunsaturated fatty acids (PUFAs) revealed no consistent pattern. Total FAs (and SFAs and MUFAs) in all species showed significant negative correlations with N cell quota (Q_N) under N deficiency, but PUFAs had species‐specific correlations with Q_N. The results show that characteristic FA profiles of algal genus or species (representing particular algal classes) underlie fluctuations according to culture conditions. The significant correlation between FAs and Q_N under N deficiency suggests that elemental and biochemical limitation of phytoplankton should be considered mutually as determinants of food quality for zooplankton in marine ecosystems.     

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.     

N、P施肥对塔里木河上游绿洲棉花C、N、P生态化学计量特征的影响

施肥通过外源物质的添加直接干预了农田生态系统中作物元素的运移循环过程。通过野外N、P施肥试验,测定棉花各生育期碳(C)、氮(N)、磷(P)元素含量及其生物量,分析棉株C、N、P元素的分配规律,探讨棉株对生长速率调控的内在机制,获得棉株体内N、P元素的内稳性指数,并判断其限制性元素类型。结果表明:棉花C、N、P元素平均含量分别为388.7、20.97、3.43 g/kg;棉花比生长速率与N∶P、C∶P间均存在负相关关系,棉花生长符合生长速率假说;N、P元素内稳性指数H分别在1.02—5.28、1.01—4.55范围内。叶片N∶P可表征植物限制性元素类型,棉花最大生长速率所对应的叶片N∶P为13,是判断限制元素的标准;综合棉花生长速率和内稳性指数研究可知研究区棉花生长受到N、P元素的共同限制,同时,在生长前期更易受P元素的限制,生长后期更易受N元素的限制。     

<Emphasis Type="Italic">Daphnia</Emphasis> growth rates in arctic ponds: limitation by nutrients or carbon?

Arctic organisms with annual life cycles experience a strong selective pressure to fulfill their life cycle at low temperatures within a short seasonal window. Yet, apart from low temperature, the factors that constrain or promote growth rates in high arctic systems are still poorly understood. A substantial part of the freshwater systems in the arctic consist of shallow, fish-free ponds with the crustacean Daphnia as the key grazer. This grazer has high demands for phosphorus (P) for RNA-synthesis and subsequently protein synthesis for growth. In this study, we compared growth of juvenile Daphnia that were fed seston from two high-Arctic (79°N) ponds on Svalbard in 2004, which differed strongly in P-content and C:P-ratios. In both ponds, Daphnia growth was limited by food quantity (carbon) rather than by P or N. The study also suggests that in absence of predators, infection level of epibionts might be an important factor regulating growth rate and population development of Daphnia growth in these systems.