Experimental demonstration of the roles of bacteria and bacterivorous protozoa in plankton nutrient cycles

We have used a model food chain composed of a natural bacterial assemblage, a pennate diatom and a bacterivorous microflagellate to investigate the factors controlling the relative importance of bacteria and protozoa as sources for regenerated nitrogen in plankton communities. In bacterized diatom cultures in which diatom growth was nitrogen-limited, the carbon:nitrogen (C:N) ratio of the bacterial substrate greatly affected which population was responsible for the uptake of nitrogen. When nitrogen was added as NH ₄ ⁺ and the cultures were supplemented with glucose, the bacteria competed successfully with the algae for NH ₄ ⁺ and prevented the growth of algae by rapidly assimilating all NH ₄ ⁺ in the cultures. Bacterivorous protozoa inoculated into these cultures grazed the bacterial population and remineralized NH ₄ ⁺ , thus relieving the nitrogen limitation of algal growth and allowing an increase in algal biomass. In contrast, bacteria in cultures supplemented with the amino acid glycine (C:N = 2) were major remineralizers of nitrogen, and the influence of protozoan grazing was minimal. We conclude that the relative importance of bacteria and protozoa as nutrient regenerators in the detrital food loop is dependent largely on the overall carbon:nutrient ratio of the bacterial substrate. The role of bacterivorous protozoa as remineralizers of a growth-limiting nutrient is maximal in situations where the carbon:nutrient ratio of the bacterial substrate is high.     

Modelling interactions between phytoplankton and bacteria under nutrient-regenerating conditions

The interactions of phytoplankton and bacteria in a nitrogen-limitedsteady-state system with an organic nitrogen compound or ammoniumas the sole nitrogen source were modelled. The effects of variousalgal excretion rates and two different mathematical representationsof excretion were examined. The model predicted that higherexcretion elevated the bacterial steady-state biomass, and loweredthe algal biomass. Bacterial respiration, which directly determinednitrogen regeneration, had an important effect on the system.The bacterial growth yield in the model was mainly a functionof the growth rate, and not of the nitrogen:carbon ratio ofthe substrate. In one version of the model, where the excretionof organic carbon increased with decreasing growth rate, themodel started to oscillate when the multiplication product ofmaximum specific excretion of excreted organic carbon (EOC)and the bacterial yield on EOC exceeded the dilution rate, irrespectiveof the form of nitrogen (ammonium or dissolved organic nitrogen)in the medium. The model results were compared with chemostatexperiments with the alga Emiliania huxleyi and a bacterialisolate in pure and mixed culture at two different dilutionrates. The carbon and nitrogen biomass of the bacteria was {smalltilde}1.5 times higher in mixed culture than in pure culture.In the experiments with low dilution rate, the recovery of nitrogenin the form of biomass, ammonium or amino acids was low, suggestingthe excretion by the algae of a refractory nitrogen-containingproduct which the bacteria could not use.     

Model of copepod growth influenced by the food carbon:nitrogen ratio and concentration, under the hypothesis of strict homeostasis

This study describes a model which addresses the processes of ingestion, assimilation, respiration, excretion and growth of copepods as a function of the concentration of food and its elemental composition in terms of carbon and nitrogen (N). Two experimental data sets are used to estimate several parameters of the model concerned with the influence of food quality. The results of the model suggest that the concentration of food and its quality (i.e. the C:N ratio) largely determine copepod growth. Both the experimental data sets and the model output show that low carbon relative to the nitrogen content of food does not limit the production of copepods. Comparing the results of the model to those of a previous model on bacteria suggests large differences between bacterial and copepod physiological responses to a variable quality of the substrate or food. The results of these models suggest that the regeneration of ammonium performed by copepods always favors regenerated primary production, whereas that performed by bacteria, depending on the quality of assimilated substrates, can favor or limit regenerated production.      

Seasonal variability in feeding behaviour, metabolic rates and carbon and nitrogen balances in the Sydney oyster, Saccostrea glomerata (Gould)

To establish if nutrients limit the growth of bivalves requires information not only on the quality of food available, but also the animals' feeding behaviour and endogenous metabolic demands. We hypothesized that growth of the Sydney rock oyster (Saccostrea glomerata) would vary in response to seasonal changes in food quality rather than quantity. We also predicted that the oysters would show feeding preferences for nitrogen over carbon, and this behaviour would result in carbon/nitrogen (C/N) ratios for ingested and absorbed matter that would be lower than the C/N of both the seston and the oysters' estimated metabolic maintenance requirements. The experiments were done in two phases under natural conditions. In phase 1, feeding behaviour was assessed on a single occasion and the results used to pose hypotheses for testing in phase 2, which included measurements made on three occasions encompassing autumn, winter and spring conditions. Growth rates varied with changes in ambient food quality and not with the concentration of total suspended matter. Feeding behaviour responded to food quality and, in most cases, resulted in nitrogen enrichment. For example, when nitrogen was potentially limiting to growth and/or maintenance, due to high food C/N (July) or high nitrogen demand (March), pre-ingestive selection ensured nitrogen enrichment of ingested matter and C/N ratios of ingested matter which were below the maintenance requirement. However, in November, when endogenous demands indicated an increased requirement for carbon, feeding behaviour resulted in carbon enrichment, an increase in carbon conversion efficiency, and ingested C/N ratios greater than the maintenance requirement. The results support the assertion of variable feeding physiology in oysters, responsive to both exogenous (seasonal differences in carbon and nitrogen availability) and endogenous (cycles of reproduction and growth) factors.     

Stoichiometry and food-chain dynamics

Traditional models of chemostat systems looking at interactions between predator, prey and nutrients have used only a single currency, such as energy or nitrogen. In reality, growth of autotrophs and heterotrophs may be limited by various elements, e.g. carbon, nitrogen, phosphorous or iron. In this study we develop a dynamic energy budget model chemostat which has both carbon and nitrogen as currencies, and examine how the dual availability of these elements affects the growth of phytoplankton, trophic transfer to zooplankton, and the resulting stability of the chemostat ecosystem. Both species have two reserve pools to obtain a larger metabolic flexibility with respect to changing external environments. Mineral nitrogen and carbon form the base of the food chain, and they are supplied at a constant rate. In addition, the biota in the chemostat recycle nutrients by means of respiration and excretion, and organic detritus is recycled at a fixed rate. We use numerical bifurcation analysis to assess the model's dynamic behavior. In the model, phytoplankton is nitrogen limited, and nitrogen enrichment can lead to oscillations and multiple stable states. Moreover, we found that recycling has a destabilizing effect on the food chain due to the increased repletion of mineral nutrients. We found that both carbon and nitrogen enrichment stimulate zooplankton growth. Therefore, we conclude that the concept of single-element limitation may not be applicable in an ecosystem context.     

Carbon and nitrogen relationships in the feeding and growth of the Pacific oyster, Crassostrea gigas (Thunberg)

Bivalve molluscs, in common with consumers in general, use behavioral and physiological mechanisms to balance metabolic requirements with available nutrients. This study considered how the Pacific oyster, Crassostrea gigas, meets the demands of growth and maintenance, measured in terms of carbon and nitrogen, in a variable food environment. Stoichiometry theory helped to evaluate: a) whether feeding behaviour modifies the intake of C and N given seasonal variability in food quality: b) how rates of metabolism and excretion, and C and N growth efficiencies, respond to mismatch between nutrient intake and the oysters' needs. Two field experiments in the Port Stephens estuary, near Sydney, Australia, measured feeding behaviour, metabolic and growth rates relative to seasonal changes in food supply. In a laboratory experiment, relationships between physiological rates and growth were measured to test a model of growth as a function of absorption of C and N. Potential metabolic targets for compensation were the C/N ratios of body tissues, maintenance and/or of soft tissue added as growth. C/N of whole soft issues varied little during the year (mean 5.4). In July (a time of low food availability of poor quality) growth was negligible and the C/N (maintenance) target was 6.7. In March (abundant food of high quality) growth was rapid with a high N-demand; the C/N of growth was 3.9. In November (medium food quality) there was an enhanced C-demand for glycogen storage; the C/N of growth was 7.9. Feeding behaviour changed the balance between C and N intake across months, primarily due to changes in the selection efficiency for nitrogen, which was highest at low filtration rates on particles of high C/N ratio. Nitrogen intake was favoured over C in July. In November, C-intake increased relative to N. In March, when abundant food nitrogen coincided with a high demand for growth, feeding behaviour was neutral with respect to C/N ratios. In all cases C/N of absorbed matter was greater than the C/N of growth. Growth efficiencies for carbon declined with increased C/N of ingested matter due to higher metabolic increments (SDA) when feeding on lower food quality; the metabolic costs of growth did not vary. In contrast, growth efficiencies for nitrogen did not alter with C/N for ingested matter, due in part to increased nitrogen losses, relative to tissue nitrogen content, when feeding on low C/N food. Nitrogen was therefore conserved metabolically relative to C. Both feeding and metabolic processes contributed to compensation for the mismatch between seasonally variable food quality and the demands of growth.     

Copepod nauplii use phosphorus from bacteria,creating a short circuit in the microbial loop

In subtropical oceans phytoplankton carbon: phosphorus (C : P) ratios are high, and these ratios are predicted to increase further with rising ocean temperatures and stratification. Prey stoichiometry may pose a problem for copepod zooplankton nauplii, which have high phosphorus demands due to rapid growth. We hypothesised that nauplii meet this demand by consuming bacteria. Naupliar bacterial and phytoplankton carbon and phosphorus ingestion, assimilation and incorporation were traced using ³³P and ¹⁴C radioisotopes. Bacterial carbon was incorporated four times less efficiently into biomass than phytoplankton carbon. In contrast, bacterial and phytoplankton phosphorus were incorporated at similar efficiencies, and bacteria could meet a substantial amount of naupliar phosphorus requirements. As parts of the ocean become more oligotrophic, bacteria could help sustain naupliar growth and survival under suboptimal stoichiometric conditions. Thus, nauplii may be a shortcut for phosphorus from the microbial loop to the classical food web.     

Stable nitrogen and carbon isotope ratios in wild native honeybees: the influence of land use and climate

The eastern hive bee Apis cerana is a major honeybee species in Asia providing numerous ecosystem services. Understanding how much the honeybees depend on natural and human-influenced plants and landscapes in different climates is important could contribute to evaluate how wild honeybees use food resources and to measure the ecosystem services. We investigated the effects of land use and climate changes on stable nitrogen and carbon isotope ratios in wild populations of A. cerana. In populations from 139 individual sites throughout Japan, we measured nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotope ratios and analyzed the effects of land use and climate. Our results showed that forested areas and annual precipitation had significant effects on δ¹⁵N, and that paddy fields and urban areas had significant effects on δ¹³C. These results suggest that A. cerana sensibly uses available food resources in the various environments and that stable nitrogen and carbon isotope ratios clearly reflect the effects of land use and climate changes on the populations of A. cerana. Thus, stable nitrogen and carbon isotope ratios in A. cerana, which widely distributes in Asia, can be used as indicators of the environments, such as land use and climate, of an area within its foraging range.     

Growth Phase and Elemental Stoichiometry of Bacterial Prey Influences Ciliate Grazing Selectivity

ABSTRACT. Protozoa are known to selectively graze bacteria and can differentiate prey based on size and viability, but less is known about the effects of prey cellular composition on predator selectivity. We measured the effect of growth phase and elemental stoichiometry of Escherichia coli on grazing by two ciliates, Euplotes vannus and Cyclidium glaucoma . Bacterial cells of a single strain were transformed with green and red fluorescent protein and harvested from culture at differing growth stages. Cells in exponential growth phase had low carbon:phosphorus (39) and nitrogen:phosphorus (9) ratios, while cells from stationary phase had high carbon:phosphorus of 104 and nitrogen:phosphorus of 26. When offered an equal mixture of both types of bacteria, Cyclidium grazed stationary phase, high carbon:phosphorus, high nitrogen:phosphorus cells to 22% of initial abundance within 135 min, while Euplotes reduced these cells to 33%. Neither ciliate species decreased the abundance of the exponential phase cells, lower carbon:phosphorus and nitrogen:phosphorus, relative to control treatments. Because protozoa have higher nitrogen:phosphorus and carbon:phosphorus ratios than their prokaryotic prey, this study raises the possibility that it may be advantageous for protozoa to preferentially consume more slowly growing bacteria.     

Growth and survival of Daphnia in epilimnetic and metalimnetic water from oligotrophic lakes: the effects of food and temperature

SUMMARY 1. In oligotrophic lakes, phytoplankton and bacteria growing in the deep chlorophyll maximum in the cool metalimnion of lakes often dominate biomass and production, but the importance of this source of food for zooplankton is unknown.
2. During much of the day, Daphnia rosea in two mountain lakes inhabited deep chlorophyll layers where food availability was at least equal to that in the epilimnion.
3. To determine the importance of the two strata (epilimnion and metalimnion) for Daphnia , we used a cross-classified factorial experiment to measure how epilimnetic and metalimnetic food and temperature (10 and 16 °C) influenced survival, growth and reproduction.
4. Daphnia survived and grew better when fed seston from the epilimnion of one lake, although chlorophyll, particulate nitrogen and particulate carbon were 2–2.5 times greater in the metalimnion.
5. Temperature had no significant influence on Daphnia survival or growth. Similar results were obtained with food from the second lake, with Daphnia surviving and reproducing better when provided with epilimnetic, rather than metalimnetic food, although the quantities of chlorophyll and carbon in the two strata were similar.
6. Food quality, rather than quantity or temperature, appeared to be the most important determinant influencing survival, growth and reproduction, and the greater food quantity in the metalimnia was not used effectively by the Daphnia .     

毛竹高速生长期土壤碳氮动态及其微生物特性

研究了典型毛竹林毛竹高速生长期间土壤碳氮动态及其微生物生态特性。结果表明:毛竹高速生长期间,3个试验地土壤全氮、碱解氮、铵态氮、硝态氮及总有机碳和水溶性有机碳(DOC)的含量均有不同幅度的下降,其中25℃蒸馏水提取DOC(25℃DOC)降幅分别达到51%、22%和223%,且25℃DOC下降幅度明显大于80℃DOC的下降幅度。随毛竹生长,土壤全氮和有机碳含量变化较为明显,相关分析表明两者呈极显著的正相关(R2=0.89**)。同时,土壤微生物量碳含量大幅度降低,由原来的800 mg/kg降到了525 mg/kg。采用PLFA法对土壤微生物群落结构进行了分析,代表细菌的饱和脂肪酸(14:0,16:0,18:0,20:0,i15:0,i16:0,i17:0,i18:0,a15:0,a17:0)基本上都分布在载荷图的右侧;代表真菌的不饱和脂肪酸(18:2w6,9c/18:0ANTE)分布在主成分载荷图的左侧,表明随着毛竹生长,土壤中细菌含量减少,真菌含量增加。说明毛竹的高速生长消耗了土壤中的碳氮,同时对土壤微生物群落结构产生了明显的影响。     

Denitrifiers associated with methanotrophs and their potential impact on the nitrogen cycle

Here I describe how losses of fixed nitrogen can occur in riparian zones by the activity of denitrifying bacteria associated with methane-oxidizing (methanotrophic) bacteria. Several methanotrophs catalyze nitrogen cycle processes that can occur in riparian buffer zones, including nitrification and nitrogen fixation. Methanotrophs can produce nitric and nitrous oxides during oxidation of ammonium (nitrification), but they cannot carry out denitrification. However, there is good evidence that denitrifying bacteria can be associated with methanotrophs and can use simple carbon compounds released by the methanotrophs as substrates for the denitrification reactions and for growth. Evidence is presented that denitrifiers isolated from methanotrophic gel-stabilized oxygen gradient systems can use methanol, formaldehyde, and formate, all methane oxidation intermediates, to support their denitrification. Such denitrification associated with methanotrophs can release dinitrogen and so contributes to losses of fixed nitrogen, and may also produce the important atmospheric trace gases nitric and nitrous oxides. Data presented also show that some methanotrophs produce nitrogen oxides, including nitrite, nitric oxide, and nitrous oxide, during growth on nitrate. Assimilatory reduction of nitrate appears to be a requirement for the release of these products.     

浮游动物稳定碳、氮同位素特征及其在水生态系统研究中的应用

浮游动物在水域生态系统食物网中起着承上启下的作用,其稳定碳、氮同位素(δ¹³C、δ¹⁵N)被广泛用于研究对食物资源的利用,揭示物质循环路径及与其他生物之间的营养关系等.浮游动物的稳定同位素也常用于解释初级生产力,判断水体污染物和环境变化过程.明确浮游动物稳定同位素组成的时空变化及影响因子,对于以其稳定同位素组成作为研究水生态系统生态过程和环境变化的指示物尤为重要.本综述基于国内外学者对水生态系统中浮游动物δ¹³C、δ¹⁵N特征及其应用的研究成果,梳理了浮游动物δ¹³C 和δ¹⁵N在不同类型水体、不同类群及时空变化的差异:浮游动物δ¹³C和δ¹⁵N时空变化则主要受食物来源及其同位素组成的影响,后者又受到初级生产力和氮源及水体物理化学性质等因素的影响.未来,浮游动物稳定同位素技术通过与新兴污染物、新兴技术及统计分析方法结合,将在地学、生物学等领域形成多手段、多学科、多尺度的研究方向.     

Growth dynamics of Azospirillum lipoferum at steady and transitory states in the presence of NH

AIMS: The objective of this paper was to study the adaptation dynamics and biochemical response of Azospirillum lipoferum grown in a continuous culture at various environmental shifts. METHODS AND RESULTS: The kinetics of A. lipoferum Sp 59b grown at steady states in a microaerobic chemostatic environment deviated from a typical Monod kinetics in both low and high dilution rates (D) due to several metabolic shifts that occurred in the microbial cell. When NH4Cl was exhausted (at low D), the microbial cell partitioned carbon flow in order to sustain growth, nitrogen fixation and assimilation processes (occurred via the glutamate synthase reaction). Increasing D the specific activities of the enzymes involved in the tricarboxylic acid cycle and the respiration rate were decreased. At transitory states, under optimal for nitrogen fixation dissolved oxygen (DO) concentrations, ammonium nitrogen negatively affected, besides nitrogen fixing activity, the bacterial growth. At sub-optimal for nitrogen fixation DO concentration (i.e. 1.56 microM) and 0.1 g l(-1) NH4Cl in the fed medium, the activities of citrate synthase and succinate dehydrogenase were significantly reduced. CONCLUSIONS: Important shifts in both carbon and nitrogen metabolism occur in A. lipoferum grown in the presence of the ammonium nitrogen, while the boundaries of ammonium nitrogen concentration in which A. lipoferum can be adapted depend on the DO concentration in the growth environment. SIGNIFICANCE AND IMPACT OF THE STUDY: Studies on growth dynamics and physiology of A. lipoferum, grown in experimental model systems, can contribute to an efficient application of these bacteria as plant-growth-promoting-agents.     

Development of a constructed subsurface-flow wetland simulation model

This paper presents a mechanistic, compartmental simulation model of subsurface-flow constructed wetlands. The model consists of six submodels, including the nitrogen and carbon cycles, both autotrophic and heterotrophic bacteria growth and metabolism, and water and oxygen balances. Data from an existing constructed wetland in Maryland were used to calibrate the model. Model results reproduced seasonal trends well. Interactions between the carbon, nitrogen, and oxygen cycles were evident in model output. In general, effluent biochemical oxygen demand, organic nitrogen, ammonium and nitrate concentrations were predicted well. Because little is known about rootzone aeration by wetland plants, oxygen predictions were fair. The model is generally insensitive to changes in individual parameters. This is due to the complexity of the ecosystem and the model, as well as the numerous feedback mechanisms. The model is most sensitive to changes in parameters that affect microbial growth and substrate use directly. This dynamic, compartmental, simulation model is an effective tool for evaluating the performance of subsurface-flow constructed wetlands. The model provided insights into treatment problems at an existing constructed wetland. With further evaluation and refinement, the model will be a useful design tool for subsurface-flow constructed wetlands.     

An Insect Herbivore Microbiome with High Plant Biomass-Degrading Capacity

Herbivores can gain indirect access to recalcitrant carbon present in plant cell walls through symbiotic associations with lignocellulolytic microbes. A paradigmatic example is the leaf-cutter ant (Tribe: Attini), which uses fresh leaves to cultivate a fungus for food in specialized gardens. Using a combination of sugar composition analyses, metagenomics, and whole-genome sequencing, we reveal that the fungus garden microbiome of leaf-cutter ants is composed of a diverse community of bacteria with high plant biomass-degrading capacity. Comparison of this microbiome''s predicted carbohydrate-degrading enzyme profile with other metagenomes shows closest similarity to the bovine rumen, indicating evolutionary convergence of plant biomass degrading potential between two important herbivorous animals. Genomic and physiological characterization of two dominant bacteria in the fungus garden microbiome provides evidence of their capacity to degrade cellulose. Given the recent interest in cellulosic biofuels, understanding how large-scale and rapid plant biomass degradation occurs in a highly evolved insect herbivore is of particular relevance for bioenergy.     

Biotransformation of phosphogypsum in media containing different forms of nitrogen

Studies on the biotransformation of phosphogypsum (a waste product formed in the course of the production of phosphorous fertilizers) with the use of sulfate reducing bacteria (SRB) demonstrated that it is a good source of sulfates and biogenic elements for these bacteria, though the addition of organic carbon and nitrogen is necessary. The aim of this study was to investigate the form of nitrogen and C:N ratio in the medium on the growth of SRB community in cultures containing phosphogypsum. Batch community cultures of sulfate reducing bacteria were maintained in medium with phosphogypsum (5.0 g/l), different concentrations of sodium lactate (1.6 - 9.4 g/l) and different forms (NH4CI, CO(NH2)2, KNO3) and concentrations (0 - 250 mg/l) of nitrogen. The growth of SRB was studied in the C:N ratio of from 2:1 to 300:1. It was found that: 1 - the best source of nitrogen for SRB is urea, followed by ammonium, the worst were nitrates; 2 - the bacteria were also able to grow in medium without nitrogen but their activity was then by approximately 15% lower than in optimal growth conditions; 3 - in medium with KNO3 inhibition of sulfate reduction by approx. 50% was observed; 4 - the highest reduction of nitrates (removal of nitrate) in media with phosphogypsum and nitrates was at limiting concentrations of sodium lactate. This is probably caused by the selection under these conditions (low concentration of hydrogen sulfide) of denitrifying bacteria or sulfate reducing bacteria capable of using nitrates as an electron acceptor.     

Nitrogen Mineralization by Acanthamoeba polyphaga in Grazed Pseudomonas paucimobilis Populations

Nitrogen mineralization was studied in a simple grazing system in which the protozoan Acanthamoeba polyphaga was grown with the bacterium Pseudomonas paucimobilis (two soil organisms isolated from the shortgrass prairie in northern Colorado). In different experiments, either carbon or nitrogen was adjusted to be in limiting amounts. When carbon was limiting, grazers were almost entirely responsible for nitrogen mineralization, with bacteria themselves contributing little. When nitrogen was limiting, nitrogen mineralization by grazers permitted continued growth by the grazed bacteria and a greater bacterial biomass production. The increased growth of the grazed bacteria did not result in an increased total amount of carbon used, but the grazed bacteria used carbon more efficiently than the ungrazed bacteria.     

Effect of nitrogen limitation and nature of the feed upon Oenococcus oeni metabolism and extracellular protein production

AIMS: The aim of the study was to characterize the effect of various nitrogen sources on Oenococcus oeni growth, carbon source utilization, extracellular protease activity and extracellular proteins. More generally, the goal is to understand how nitrogen-based additives might act to enhance malolactic fermentation in wine. METHODS AND RESULTS: Five yeast extracts were used. As the amino acid and nitrogen analyses revealed, they were similar in global amino acid composition, except for arginine level. Nevertheless the ratio of amino acids between free/bound, and low/high molecular weight fractions were highly different. One of the yeast extracts led to a significant protease activity in the supernatant and to a poor final biomass of the IOB84.13 strain compared to the other ones. For the IOB84.13 strain specifically, arginine addition to the arginine poor yeast extract did not restore growth. 35S-methionine-labelled extracellular proteins were separated by SDS-PAGE. Signals were detected in all media early in the growth phase and were maintained during 48 h of culture. CONCLUSIONS: A significant protease activity was detected for O. oeni supernatants during growth under nitrogen limitation but only for certain nitrogen sources. Moreover, the activity was strain dependent. Peptides (0.5-10 kDa) seemed to be more favourable for growth of wine bacteria than <0.5 kDa nitrogen sources. The extracellular protein signal patterns differed more greatly between the bacterial strains tested than between the nitrogen molecules in the medium. SIGNIFICANCE AND IMPACT OF THE STUDY: This is the first study extensively considering the role of the nitrogen source composition and level upon O. oeni growth and metabolism.     

The Influence of Carbon Dioxide and Daily Photon-flux Density on Optimal Leaf Nitrogen Concentration and Root: Shoot Ratio

Using a cost-benefit model, the leaf nitrogen concentrationand root : shoot ratio that maximize whole-plant relative growthrate are determined as a function of the above-ground environment(integrated daily photon flux density and the concentrationof carbon dioxide at the site of fixation within the leaf).The major advantage of this approach is that it determines theadaptive significance of leaf physiology by considering thefunctional integration of leaves and roots. The predicted responseto increasing daily photon flux densities is an increase inoptimal leaf N concentration (N_opt) and a concomitant increasein root: shoot ratio. Increased carbon dioxide concentrations,on the other hand, reduce N_opt and only slightly change root:shoot ratio. The observed increase in leaf nitrogen concentrationfound in plants growing at high altitudes (low CO₂ partial pressure)is also predicted. Since these responses to light and CO₂ maximizethe whole-plant relative growth rate, the observed adjustmentsthat plants make to light and carbon dioxide concentration appearto be adaptive. We show that the relationship between photosynthesis and leafnitrogen concentration is complex and depends on the light andCO₂ levels at which photosynthesis is measured. The shape ofthis function is important in determining N_opt and the oppositeresponse of leaf nitrogen to light and carbon dioxide is shownto be the result of the different effects of light and CO₂ onthe photosynthesis-leaf nitrogen curve. Plant growth, photosynthesis, leaf nitrogen, biomass allocation, optimization, carbon dioxide light