The light : nutrient ratio in lakes: a test of hypothesized trends in bacterial nutrient limitation

The light-to-nutrient hypothesis explores how the balance between energy (as light energy) and nutrients (as total phosphorus) shapes aquatic ecosystem structure and process. The balance of energy and nutrients is thought to regulate ecosystem structure and process such that, in a "high" light-to-nutrient environment, bacteria would probably be driven towards phosphorus (P) limitation, whereas, in a "low" light-to-nutrient environment, bacteria would be driven towards carbon (C) limitation. We assessed the growth limitation of bacteria in two reservoirs of the southern U.S.A. using a mortality-corrected dilution-growth approach. We compared the frequency of P and C growth limitation with the intralake variation in the light-to-nutrient environment. As a metric of the light-to-nutrient environment, we used the ratio of the mean light in the surface mixed layer ( I _m) to the total phosphorus concentration ( TP ). In each lake, bacterial growth was more often P-limited when the I _m :  TP ratio was above the median ratio than below. We believe our data provide the first evidence supporting this aspect of the light-to-nutrient hypothesis.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Influence of nutrient availability on phytoplankton growth and community structure in the Port Adelaide River,Australia: [2pt] bioassay assessment of potential nutrient limitation

We investigated the influence of nutrient availability, specifically nitrogen, phosphorus and silicon on growth and community structure of phytoplankton from the Port Adelaide River estuary, South Australia. Two bioassay experiments were conducted. The first, Nutrich1, involved addition of nutrients in vitro to samples of the natural phytoplankton community from a single location in the upper estuary. The second, Nutrich2, involved nutrient addition and incubation of water from five locations in the estuary following inoculation with a `standardised' phytoplankton assemblage derived from laboratory cultures. In Nutrich1, enrichment with silicon led to greatly enhanced phytoplankton biomass due to increased growth of diatoms. Addition of nitrogen or phosphorus had little effect on phytoplankton growth. In Nutrich2, addition of nitrogen resulted in enhanced growth of phytoplankton in water collected from near the mouth the estuary, but there were no differences in growth among nutrient treatments for the remaining locations. Comparison of phytoplankton growth rate among locations revealed a trend of decreasing growth in moving towards the mouth of the estuary. This trend was unaffected by enrichment with nitrate, phosphate or silicate. We suggest that spatial variation in growth potential within the Port Adelaide River estuary may relate to variation in the concentration of nitrogen as ammonium.     

Relative importance of competition with Daphnia (Cladocera) and nutrient limitation on Anuraeopsis (Rotifera) population dynamics in a laboratory study

• 1 In the laboratory, the growth and reproduction of Anuraeopsis fissa were measured when fed on Scenedesmus species grown in nutrient‐sufficient, nitrogen‐limited and phosphorus‐limited media and in the presence or absence of one adult Daphnia longispina per vial.
• 2 Poor food quality may reduce the effect of competition on rotifers. Competition from Daphnia was stronger with nutrient sufficient algae than with nutrient limited algae. P‐limitation significantly reduced Anuraeopsis population growth rate and fecundity. The effect of nutrient limitation on Anuraeopsis was stronger than that of competition with Daphnia. The Anuraeopsis population declined with P‐limited food in both the presence and absence of Daphnia.
• 3 Exploitative competition by Daphnia on Anuraeopsis was stronger in the nutrient‐sufficient treatment than in the N‐limited one. Density, fecundity and population growth rate of Anuraeopsis were negatively affected by Daphnia in the nutrient‐sufficient treatment, while only fecundity was reduced by Daphnia in the N‐limited treatment. Consequently, in the N‐limited treatment, mortality should be lower in the presence of Daphnia. This result could suggest that Anuraeopsis lives longer when short of nitrogen.
• 4 Nutrient limitation may affect to the competitive interactions between zooplankton species. P‐limitation decreased the quality of algae as food for Anuraeopsis while N‐limitation decreased the susceptibility of this rotifer species to exploitative competition by Daphnia.

     

Control of cell morphology of the yeast Saccharomyces cerevisiae by nutrient limitation in continuous culture

Saccharomyces cerevisiae IFO 0203, a polyploid yeast used in ethanol production in Japan, grows as ovoid cells in unstirred batch culture and on fully nutritive agar plates (2% w/v glucose; 0.67% w/v Difco yeast nitrogen base).
Extensively branched pseudohyphae formed on 0.01% w/v ammonium sulphate plates within a few days. In continuous culture with high oxygen supply and limiting glucose, cells were elongated but growth was vigorous and the daughter cells separated well after budding.
Limitation of growth by either nitrogen source or oxygen during continuous culture resulted in formation of truncated, occasionally branched, pseudohyphae up to five cells in length.     

Flocculation onset in Saccharomyces cerevisiae: the role of nutrients

AIMS: To examine the role of the nutrients on the onset of flocculation in an ale-brewing strain, Saccharomyces cerevisiae NCYC 1195. METHODS AND RESULTS: Flocculation was evaluated using the method of Soares, E.V. and Vroman, A. [Journal of Applied Microbiology (2003) 95, 325]. For cells grown in chemically defined medium (yeast nitrogen base with glucose) or in rich medium (containing yeast extract, peptone and fermentable sugars: fructose or maltose), the onset of flocculation occurred after the end of exponential respiro-fermentative phase of growth being coincident with the attainment of the lower level of carbon source in the culture medium. Cells, in exponential respiro-fermentative phase of growth, transferred to a glucose-containing medium without nitrogen source, developed a flocculent phenotype, while these carbon source starved cells, in the presence of all other nutrients that support growth, did not flocculate. In addition, cells in exponential phase of growth, under catabolite repression, when transferred to a medium containing 0.2% (w/v) of fermentable sugar (fructose or maltose) or 2% (v/v) ethanol, showed a rapid triggering of flocculation, while when incubated in 2% (v/v) glycerol did not develop a flocculent phenotype. CONCLUSIONS: The onset of flocculation occurs when a low sugar and/or nitrogen concentration is reached in culture media. The triggering of flocculation is an energetic dependent process influenced by the carbon source metabolism. The presence of external nitrogen source is not necessary for developing a flocculent phenotype. SIGNIFICANCE AND IMPACT OF THE STUDY: This work contributes to the elucidation of the role of nutrients on the onset of flocculation in NewFlo phenotype yeast strains. This information might be useful to the brewing industry, in the control of yeast flocculation, as the time when the onset of flocculation occurs can determine the fermentation performance and the beer quality.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.     

Role of nutrient limitation in the competition between uropathogenic strains of Klebsiella pneumoniae and Escherichia coli in mixed biofilms

Klebsiella pneumoniae and Escherichia coli form mixed species biofilms in catheter-associated urinary tract infections. Recently, a detrimental effect of K. pneumoniae over E. coli was observed in mixed species biofilms grown in an artificial urine medium. The mechanism behind this competitive interaction was studied. K. pneumoniae partially outcompeted E. coli in early-stage batch-fed biofilms, whereas both microorganisms co-exist at longer times (K. pneumoniae:E. coli ratio, 55:1), as shown by cell counts and confocal microscopy. E. coli cells were scattered along the K. pneumoniae biofilm. Biofilm supernatants did not appear to contain either antimicrobial or anti-biofilm activities against E. coli. Biofilms grown under continuous flow prevented interspecies competition. K. pneumoniae showed both increased siderophore production and better growth in iron-limited media compared to E. coli. In summary, these results indicate the importance of nutrient (particularly iron) competition in the modulation of the bacterial composition of mixed species biofilms formed by uropathogenic K. pneumoniae and E. coli.     

Saccharomyces cerevisiae: the effect of different forms and concentrations of iodine on uptake and yeast growth

The essentiality of iodine for humans, especially in the early stages of life, is well recognized. The chemical forms of iodine in food supplements, infant formulae and iodated salt are either iodide (KI) or iodate (KIO₃). Because there are no or rare data about iodine uptake by yeasts, we investigated the influence of different sources of iodine, as KI, KIO₃ and periodate (KIO₄), on its uptake in and growth of the model yeast Saccharomyces cerevisiae . KIO₃ inhibited the growth of the yeast the most and already at a 400 μM initial concentration in the growth medium; the OD was reduced by 23% in comparison with the control, where no KIO₃ was added. The uptake of different iodine sources by the yeast S. cerevisiae was minimal, in total <1%. Tracer experiments with radioactive ¹³¹I added as KI showed that the yeast S. cerevisiae does not have the ability to transform KI into volatile species. We investigated the specificity of iodine uptake added as KIO₃ in the presence of Na₂SeO₄ or ZnCl₂ or K₂CrO₄ in the growth medium, and it was found that chromate had the most influence on reduction of KIO₃ uptake.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Characterisation of the type and extent of nutrient limitation in grassland vegetation using a bioassay with intact sods

An important methodological problem in plant ecology concerns the way in which the type and extent of nutrient limitation in terrestrial communities should be assessed. Conclusions on nutrient limitation have been founded mainly on soil extractions, fertiliser trials and tissue nutrient concentrations. In order to avoid some of the problems associated with these methods, we employed a special technique using intact sods which rooted both in the intact soil and in a nutrient solution, from which N, P and K were omitted stepwise. The method was applied to hay-field communities which differed in their history of fertiliser application. Four fields were compared which were not fertilised for 2, 6, 19 or 45 years, while hay making continued. This was done to restore former species-rich grassland communities. We tested the hypothesis that the increase in species diversity in these grasslands was attended by an increase in the number of limiting nutrients.We observed clear shifts in the type and extent of nutrient limitation. Fields which were recently fertilised were characterised by nitrogen and potassium limitation, while phosphorus limitation increased in importance towards the later stages of succession. In the last field (45 years unfertilised) N, P and K equally limited production at the community level. These conclusions differed from those drawn from a fertiliser trial in these same four fields, which failed to detect phosphorus limitation.It is concluded that the use of this method provides a valuable extra source of information while studying relationships between nutrient limitation and species diversity in grassland communities.     

太平洋中西部海域浮游植物营养盐的潜在限制

2009年8月至9月期间在太平洋西部N1站和中部N2站进行现场营养盐加富培养实验。结果显示:N1站,浮游植物生物量对N或者P添加都有较强的响应,其中N+P+Si组和N+P组浮游植物长势迅速,叶绿素a从初始的0.03μg/L分别达到2.12μg/L和1.83μg/L,同时P先于N和Si之前被耗尽;说明N1站为N、P共同限制,P是首要限制因子。而N2站,浮游植物生物量仅对N、P共同添加有明显响应,N先于P和Si被浮游植物消耗殆尽。利用培养过程中营养盐比值变化推断,N1站浮游植物以低于Redfield ratio(16N∶1P)吸收N和P;而N2站浮游植物以高于Redfield ratio(16N∶1P)吸收N和P。这可能解释了太平洋西部的寡营养盐海域为潜在P限制,而在太平洋中部海域则为潜在N限制。     

Heterogeneity in the nutrient limitation of different bacterioplankton groups in the Eastern Mediterranean Sea

The heterotrophic bacterial community of the Eastern Mediterranean Sea is believed to be limited by phosphorus (P) availability. This observation assumes that all bacterial groups are equally limited, something that has not been hitherto examined. To test this hypothesis, we performed nutrient addition experiments and investigated the response of probe-identified groups using microautoradiography combined with catalyzed reporter deposition fluorescence in situ hybridization. Our results show contrasting responses between the bacterial groups, with Gammaproteobacteria being the group more affected by P availability. The Roseobacter clade was likely colimited by P and nitrogen (N), whereas Bacteroidetes by P, N and organic carbon (C). In contrast, SAR11 cells were active regardless of the nutrient concentration. These results indicate that there is high heterogeneity in the nutrient limitation of the different components of the bacterioplankton community.     

Flocculation in Saccharomyces cerevisiae: a review

The present work reviews and critically discusses the aspects that influence yeast flocculation, namely the chemical characteristics of the medium (pH and the presence of bivalent ions), fermentation conditions (oxygen, sugars, growth temperature and ethanol concentration) and the expression of specific genes such as FLO1, Lg‐FLO1, FLO5, FLO8, FLO9 and FLO10. In addition, the metabolic control of loss and onset of flocculation is reviewed and updated. Flocculation has been traditionally used in brewing production as an easy and off‐cost cell‐broth separation process. The advantages of using flocculent yeast strains in the production of other alcoholic beverages (wine, cachaça and sparkling wine), in the production of renewal fuels (bio‐ethanol), in modern biotechnology (production of heterologous proteins) and in environmental applications (bioremediation of heavy metals) are highlighted. Finally, the possibility of aggregation of yeast cells in flocs, as an example of social behaviour (a communitarian strategy for long‐time survival or a means of protection against negative environmental conditions), is discussed.     

Nutrient limitation and soil development: Experimental test of a biogeochemical theory

Walker & Syers (1976) proposed a conceptual model that describesthe pattern and regulation of soil nutrient pools and availability during long-term soil and ecosystem development. Their model implies that plantproduction generally should be limited by N on young soils and by P on oldsoils; N and P supply should more or less equilibrate onintermediate-aged soils. We tested the application of this model to nutrientlimitation, using a well characterized substrate age sequence in Hawaiianmontane rain forest. Earlier experiments had evaluated nutrient limitationin forests on a young (300 y) and an old (4,100,000 y) substrate on the samedevelopmental sequence; N alone limited tree growth on the youngsubstrate, while P alone did so on the old one. An additional fertilizerexperiment based on replicated treatments with N, P, and all othernutrients combined, applied in individually and in all factorialcombinations, was established in an intermediate-aged site in theLaupahoehoe Forest Reserve, Hawaii. Here, diameter increments of thedominant tree Metrosideros polymorpha increased slightly with Nadditions, and nearly doubled when N and P were added together.Additions of elements other than N and P had no significant effecton growth. These results show that N and P had equilibrated (relativeto plant requirements) in the intermediate aged site. Together withthe earlier experiments, these results suggest that the Walker and Syersmodel provides a useful starting point for explaining the nature anddistribution of nutrient limitation in forest ecosystems.     

Variation in nutrient limitation of lotic and lentic algal communities in a Texas (USA) river

Nutrient limitation of periphyton and phytoplankton was assessed in the Upper Guadalupe River, Texas USA. Nutrient-diffusing substrates with added nitrogen (N) and phosphorus (P) were used to identify the limiting nutrient for lotic algae at three river sites in summer, fall, and winter. Pots enriched with P had significantly higher chlorophyll a concentrations for 7 of 9 trials. Added N alone did not significantly increase algal standing crops, although it was found to be secondarily limiting on one (and possibly two) occasions. Flow-through enrichment experiments were conducted in order to quantify the concentration of P needed to significantly increase algal standing crops. Response to enrichment was rapid when ambient P concentration was low (< 0.010 mg L^–1), but more moderate when ambient P levels were higher (0.015–0.025 mg L^–1). Nutrient limitation of phytoplankton in small surface-release reservoirs varied throughout the study, but N was either primarily or secondarily limiting in 6 of 8 trials; shifts in the limiting nutrient were correlated with fluctuations in flow into the reservoirs. Our enrichment studies show that algal response to nutrient addition was unpredictable as phytoplankton tended to be N-limited while periphyton was mainly P-limited. Further, while discharge apparently dictated the nutrient-biomass relationship for phytoplankton in reservoirs, ambient nutrient level is an important determinant of lotic periphyton response to enrichment.     

Strong growth limitation of a floating plant (Lemna gibba) by the submerged macrophyte (Elodea nuttallii) under laboratory conditions

1. The asymmetric competition for light and nutrients between floating and submerged aquatic plants is thought to be key in explaining why dominance by either of these groups can be stable and difficult to change. 2. Although the shading effect of floating plants on submerged plants has been well documented, the impact of submerged plants on floating plants has been poorly explored hitherto. 3. Here, we used laboratory experiments to examine how submerged plant (Elodea nuttallii) alter nutrient conditions in the water column and how this affects the growth of floating plants (Lemna gibba). 4. We demonstrate that, at higher nutrient concentrations, Lemna is increasingly likely to outcompete Elodea. 5. Under low nutrient concentrations (0.1–2 mg N L^?1) Elodea can strongly reduce the growth of Lemna. Growth of floating plants virtually stopped in some of the experiments with Elodea. 6. Extremely reduced tissue N, Mn, chlorophyll and elongated roots indicated that the growth inhibition of Lemna by Elodea was predominantly caused by the latter’s impact on the nutrient conditions for floating plants. 7. These results strengthen the hypothesis that submerged plants can prevent colonization of a lake by floating plants.     

Towards an integration of ecological stoichiometry and the metabolic theory of ecology to better understand nutrient cycling

Ecologists have long recognized that species are sustained by the flux, storage and turnover of two biological currencies: energy, which fuels biological metabolism and materials (i.e. chemical elements), which are used to construct biomass. Ecological theories often describe the dynamics of populations, communities and ecosystems in terms of either energy (e.g. population-dynamics theory) or materials (e.g. resource-competition theory). These two classes of theory have been formulated using different assumptions, and yield distinct, but often complementary predictions for the same or similar phenomena. For example, the energy-based equation of von Bertalanffy and the nutrient-based equation of Droop both describe growth. Yet, there is relatively little theoretical understanding of how these two distinct classes of theory, and the currencies they use, are interrelated. Here, we begin to address this issue by integrating models and concepts from two rapidly developing theories, the metabolic theory of ecology and ecological stoichiometry theory. We show how combining these theories, using recently published theory and data along with new theoretical formulations, leads to novel predictions on the flux, storage and turnover of energy and materials that apply to animals, plants and unicells. The theory and results presented here highlight the potential for developing a more general ecological theory that explicitly relates the energetics and stoichiometry of individuals, communities and ecosystems to subcellular structures and processes. We conclude by discussing the basic and applied implications of such a theory, and the prospects and challenges for further development.     

Co-regulation of polar mRNA transport and lifespan in budding yeast Saccharomyces cerevisiae

Recent studies have uncovered the links between aging, rejuvenation and polar protein transport in the budding yeast Saccharomyces cerevisiae. Here, we examined a still unexplored possibility for co-regulation of polar mRNA transport and lifespan. To monitor the amount and distribution of mRNA-containing granules in mother and daughter cells, we used a fluorescent mRNA-labeling system, with MFA2 as a reporter gene. The results obtained showed that deletion of the selected longevity regulators in budding yeast had a significant impact on the polar mRNA transport. This included changes in the amount of mRNA-containing granules in cytoplasm, their aggregation and distribution between the mother and daughter cells. A significant negative correlation was found between strain-specific longevity, amount of granules and total fluorescent intensity both in mother and daughter cells. As indicated by the coefficient of determination, approximately 50–75% of variation in yeast lifespan could be attributed to the differences in polar mRNA transport.