Growth rate of alpine phytoplankton assemblages from contrasting watersheds and N‐deposition regimes exposed to nitrogen and phosphorus enrichments

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Can Hydrodynamics Change Phosphorus Strategies of Diatoms?—Nutrient Levels and Diatom Blooms in Lotic and Lentic Ecosystems

Diatom blooms occur in many water bodies worldwide, causing significant ecological and social concerns. In order to understand the mechanisms of diatom blooms formation, the effects of varying phosphorus (P) concentration and hydrodynamics on the growth of diatoms were studied by combining results from field observations and laboratory experiments. The field investigation showed that spring diatom blooms (Cyclotella meneghiniana and Stephanodiscus hantzschii) occurred in Lake Taihu and Hanjiang River with similar environmental factors such as water temperature, pH, and dissolved oxygen in 2008. Concentrations of total phosphorus (TP), total nitrogen, and ammonia nitrogen (NH₄-N) in Lake Taihu were significantly higher than the concentrations in the Hanjiang River. Laboratory experiments were conducted to evaluate growth and physiological responses of four lotic diatoms (Cyclotella atomus, Fragilaria crotonensis, Nitzschia palea, and S. hantzschii, isolated from the Hanjiang River) and three lentic diatoms (C. meneghiniana, Melosira varians, and Stephanodiscus minutulus, isolated from Lake Taihu, Lake Donghu, and Guanqiao Pond, respectively) to various P concentrations under small-scale turbulent and standing conditions. Our results showed that, with turbulence, lotic diatoms C. atomus, F. crotonensis, N. palea, and S. hantzschii demonstrated a significant increase in affinity for P compared with lentic diatoms C. meneghiniana, M. varians, and S. minutulus. Moreover, lotic diatoms C. atomus, F. crotonensis, and N. palea had higher growth rates and photosynthetic efficiencies with small-scale turbulence than with standing conditions both in P-limited and P-replete conditions. Lotic species S. hantzschii and three lentic diatoms (C. meneghiniana, M. varians, and S. minutulus) grew well under standing conditions. Our results may explain our field observation that the occurrence of diatom blooms in lakes is often associated with higher TP concentrations whereas in rivers, diatom blooms occur at a wide range of TP concentrations under flows. Therefore, different hydrodynamics and nutrient concentrations determined the dominant diatom species, according to their habitat-dependent physiological characteristics.     

Erratum to: How do UV radiation, temperature, and zooplankton influence the dynamics of alpine phytoplankton communities?

Plankton in mountain lakes are confronted with generally higher levels of incident ultraviolet radiation (UVR), lower temperatures, and shorter growing seasons than their lower elevation counterparts. The direct inhibitory effects of high UVR and low temperatures on montane phytoplankton are widely recognized. Yet little is known about the indirect effects of these two abiotic factors on phytoplankton, and more specifically whether they alter zooplankton grazing rates which may in turn influence phytoplankton. Here, we report the results of field microcosm experiments that examine the impact of temperature and UVR on phytoplankton growth rates and zooplankton grazing rates (by adult female calanoid copepods). We also examine consequent changes in the absolute and relative abundance of the four dominant phytoplankton species present in the source lake (Asterionella formosa, Dinobryon sp., Discostella stelligera, and Fragilaria crotonensis). All four species exhibited higher growth rates at higher temperatures and three of the four species (all except Dinobryon) exhibited lower growth rates in the presence of UVR versus when shielded from UVR. The in situ grazing rates of zooplankton had significant effects on all species except Asterionella. Lower temperatures significantly reduced grazing rates on Fragilaria and Discostella, but not Dinobryon. While UVR had no effect on zooplankton grazing on any of the four species, there was a significant interaction effect of temperature and UVR on zooplankton grazing on Dinobryon. Discostella and Dinobryon increased in abundance relative to the other species in the presence of UVR. Colder temperatures, the presence of zooplankton, and UVR all had consistently negative effects on rates of increase in overall phytoplankton biomass. These results demonstrate the importance of indirect as well as direct effects of climate forcing by UVR and temperature on phytoplankton community composition in mountain lakes, and suggest that warmer climates and higher UVR levels may favor certain species over others.     

Nutrient limitation of phytoplankton communities in Subarctic lakes and ponds in Wapusk National Park, Canada

We determined the limiting nutrient of phytoplankton in 21 lakes and ponds in Wapusk National Park, Canada, using nutrient enrichment bioassays to assess the response of natural phytoplankton communities to nitrogen and phosphorus additions. The goal was to determine whether these Subarctic lakes and ponds were nutrient (N or P) limited, and to improve the ability to predict future impacts of increased nutrient loading associated with climate change. We found that 38% of lakes were not limited by nitrogen or phosphorus, 26% were co-limited by N and P, 26% were P-limited and 13% were N-limited. TN/TP, DIN/TP and NO₃ ⁻/TP ratios from each lake were compared to the Redfield ratio to predict the limiting nutrient; however, these predictors only agreed with 29% of the bioassay results, suggesting that nutrient ratios do not provide a true measure of nutrient limitation within this region. The N-limited lakes had significantly different phytoplankton community composition with more chrysophytes and Anabaena sp. compared to all other lakes. N and P limitation of phytoplankton communities within Wapusk National Park lakes and ponds suggests that increased phytoplankton biomass may result in response to increased nutrient loading associated with environmental change.     

Species-specific differences in phytoplankton responses to N and P enrichments and the N:P ratio in the Archipelago Sea, northern Baltic Sea

A nutrient enrichment experiment was conducted in order to studythe role of nitrogen (N), phosphorus (P) and the N:P ratio onthe early summer phytoplankton community in the ArchipelagoSea, northern Baltic Sea. The phytoplankton community was, interms of chlorophyll a and total biomass, primarily N-limited,but the individual species varied in their responses to thenutrient supply. The recorded overall N limitation was due tofast growth responses of a few N-limited species such as thediatom Chaetoceros wighamii (Brightwell) and the mixotrophicchrysophyte Uroglena sp. Another dominating diatom, Skeletonemacostatum (Greville) Cleve was most clearly P-limited. The N:Pratio had the strongest effect on Uroglena sp., which grew exponentiallyin the enrichments with a high N:P ratio. This can be explainedby the ability of the species to feed on P-rich bacteria, whichgives it a competitive advantage in P-limited conditions. Thespecies-specific differences in the responses to the nutrientenrichments can generally be explained by differences in thespecies physiology and they were consistent with the theoryof resource competition.     

Nitrogen deposition and warming – effects on phytoplankton nutrient limitation in subarctic lakes

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N‐deposition (<1 kg N ha^?1 yr^?1) in northern subarctic Sweden. Short‐term bioassay experiments with N‐ and P‐additions revealed that phytoplankton in high‐alpine lakes were more prone to P‐limitation, and with decreasing altitude became increasingly N‐ and NP‐colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N‐deposition over the Swedish mountain areas will, with the exception of high‐alpine lakes, enhance biomass and drive phytoplankton from N‐ to P‐limitation. However, if not accompanied by warming, N‐input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.     

Winter phytoplankton community structure in three shallow temperate lakes during ice cover

The general model of seasonal phytoplankton succession in temperate lakes suggests that winter phytoplankton growth is minimal under ice-cover. However, some studies have found diverse phytoplankton communities during winter. The primary objectives of this study were to determine the species composition and the changes in the winter phytoplankton community structure under the ice. For 2 consecutive winters, phytoplankton samples were collected under ice-cover at 4 sites on 3 lakes in Arrowwood National Wildlife Refuge (ANWR), near Pingree, North Dakota. Ninety taxa were identified and enumerated. Densities of several of these taxa frequently exceeded 10⁶ cells l^–1. The winter phytoplankton communities in these lakes were dominated by flagellates, principally cryptomonads, a synurophyte (Synura uvella), small chrysophytes (Chrysococcus spp., Kephyrion spp.) and a dinoflagellate (Peridinium aciculiferum), as well as non-flagellate microchlorophytes (Monoraphidium spp., Ankistrodesmus spp., and Pseudodictyosphaerium sp.), a cyanobacterium (Gloeocapsa aeruginosa) and centric diatoms (Stephanodiscus minutulus, S. parvus and Cyclotella meneghiniana).     

Relationship between phytoplankton and environmental factors in landscape water supplemented with reclaimed water

The water quality of Feng-qing Lake, which is a landscape lake supplemented with reclaimed water, was surveyed to investigate the relationship between phytoplankton and environmental variables. A total of 29 water samples were collected to analyze temporal variations of phytoplankton and environmental factors from July 2013 to June 2014. Six phyla and 39 genera of phytoplankton were identified when the lake was supplied with reclaimed water. Among these, Cyanophyta and Chlorophyta account for 38.46% and 30.77% of phytoplankton, respectively. The dominant species in the lake are Pseudanabaena limnetica and Chlorella vulgaris, which are present the entire year. Other leading species include Cosmarium sp. and Raphidiopsis curvata. Principal component analysis (PCA) was conducted to analyze the relationship among environmental factors. Canonical correspondence analysis (CCA) was performed to investigate the relationship between environment factors and dominant species. The PCA result showed that temperature (T), total phosphorus (TP), total nitrogen (TN), transparency, and dissolved oxygen are the main factors that affect the eutrophication level of the lake. The CCA result revealed that TN, PO₄³⁻–P, chemical oxygen demand (COD), T, and chlorophyll a exhibit a close relation with dominant species. In particular, TN, salinity, and COD influence the growth of P. limnetica; T and COD influence the growth of R. curvata; and T, PO₄³⁻–P, NH₃–N, and pH influence the growth of C. vulgaris and Cosmarium sp.     

Dynamics and Nutritional Ecology of a Nanoflagellate Preying Upon Bacteria

Ingestion and growth rates of the nanoflagellate predator Ochromonas danica feeding on the bacterium Pseudomonas fluorescens were quantified in laboratory cultures. Bacterial prey were grown under four nutritional conditions with respect to macronutrient elements: C-limited, N-limited, P-limited, and balanced. Ingestion and growth rates were saturating functions of prey abundance when preying upon nutritionally balanced, C-limited, and P-limited bacteria but were unimodal functions of abundance when preying on N-limited bacteria. At saturating prey concentrations, the ingestion rate of C-limited prey was about twice that of prey in other nutritional states, while at subsaturating prey concentrations, the ingestion rates of both C- and N-limited prey were higher than those of prey in other nutritional states. Over all prey concentrations, growth was most rapid on balanced and C-limited prey and generally lowest for P-limited prey. Due to the unimodal response of growth rate to abundance of N-limited prey, growth rate on N-limited prey approached that obtained on balanced and C-limited prey when prey were available at intermediate abundances. The accumulation of recycled N increased with the growth rate of O. danica. Recycling of N was highest when O. danica was feeding upon P-limited prey. The accumulation of recycled P increased with growth rate for balanced and N-limited prey, but not for P-limited prey, which consistently had low accumulation of recycled P. The low growth rate and negligible recycling of P for O. danica preying on P-limited prey is consistent with the theory of ecological stoichiometry and resembles results found for crustacean zooplankton, especially in the genus Daphnia. Potentially, the major predators of bacterioplankton and a major predator of phytoplankton play analogous roles in the trophic dynamics and biogeochemistry of aquatic ecosystems.     

Effects of Bradyrhizobium japonicum and Soybean (Glycine max (L.) Merr.) Phosphorus Nutrition on Nodulation and Dinitrogen Fixation

Cells of Bradyrhizobium japonicum were grown in media containing either 1.0 mM or 0.5 μM phosphorus. In growth pouch experiments, infection of the primary root of soybean (Glycine max (L.) Merr.) by B. japonicum USDA 31, 110, and 142 was significantly delayed when P-limited cells were applied to the root. In a greenhouse experiment, B. japonicum USDA 31, 110, 122, and 142 grown with sufficient and limiting P were used to inoculate soybeans which were grown with either 5 μM or 1 mM P nutrient solution. P-limited cells of USDA 31 and 110 formed significantly fewer nodules than did P-sufficient cells, but P-limited cells of USDA 122 and 142 formed more nodules than P-sufficient cells. The increase in nodule number by P-limited cells of USDA 142 resulted in significant increases in both nodule mass and shoot total N. In plants grown with 1 mM P, inoculation with P-limited cells of USDA 110 resulted in lower total and specific nitrogenase activities than did inoculation with P-sufficient cells. Nodule numbers, shoot dry weights, and total N and P were all higher in plants grown with 1 mM P, and plants inoculated with USDA 31 grew poorly relative to plants receiving strains USDA 110, 122, and 142. Although the effects of soybean P nutrition were more obvious than those of B. japonicum P nutrition, we feel that it is important to develop an awareness of the behavior of the bacterial symbiont under conditions of nutrient limitation similar to those found in many soils.     

Phytoplankton growth in the Ohio, Cumberland and Tennessee Rivers, USA: inter-site differences in light and nutrient limitation

Seasonal patterns in resource limitation of phytoplankton growth were assessed monthly within three large rivers with differing extents of water regulation. The Ohio River is regulated by low dams that do not substantially modify discharge, while the Cumberland and Tennessee Rivers are impounded by a series of high dams to enhance water storage for downstream flood control. Laboratory dilution assays with light and nutrient manipulations indicated that light was the main factor limiting phytoplankton growth at irradiances below 7 E m^–2 d^–1. Light limited growth was frequent in the turbid, higher discharge of the Ohio River, but was rare in the heavily regulated Tennessee and Cumberland Rivers. When irradiance exceeded 7 E m^–2 d^–1, phytoplankton were either P-limited (Cumberland River), co-limited by P and N (Tennessee River), or Si limited (Ohio River). Site-specific differences in nutrient limitation were consistent with differences in ambient nutrient levels, with the Tennessee and Cumberland Rivers characterized by lower N and P concentrations, and the Ohio River by lower Si. Downstream nutrient depletion was evident in the Ohio River through comparison of an upstream and a downstream site, with nutrient limitation (Si) occurring more frequently downstream. Phytoplankton growth rates at ambient light and nutrient levels ranged from 0.1 to 1.5 d^–1 in the Ohio River and 0.2 to 0.6 d^–1 in the Tennessee and Cumberland Rivers. Growth rates were greatest at the onset of the summer base pool, as light intensities increased and nutrient levels were maximal. Our findings indicate that multiple factors regulate phytoplankton growth in regulated rivers and that spatial complexity may arise from differences in discharge and water aging.     

Dissolved organic carbon in a humic lake: effects on bacterial production and respiration

Allochthonous matter was the main source of carbon for pelagic bacteria in a humic lake, accounting for almost 90% of the carbon required to support observed bacterial growth. The estimated contribution from zooplankton excretion was of the same magnitude as direct phytoplankton release, both accounting for 5–7% of bacterial demands for dissolved carbon. Bacteria were an important source of carbon both for heterotrophic phytoplankton and for filter feeding zooplankton species, further stressing the role of humus DOC in overall lake productivity. The high contribution of allochthonous DOC implies a stoichiometry of dissolved nutrients with a surplus of C relative to P. The high P cell quota of bacteria suggest that under such conditions they are P-limited and act like net consumers of P. Excess C will be disposed of, and bacterial respiration rate will increase following a transition from carbon-limited bacterial growth towards mineral-nutrient-limited growth. Thus the high community respiration and frequent CO₂-supersaturation in humic lakes may be caused not only by the absolute supply of organic C, but also by the stoichiometry of the dissolved nutrient pool.     

Ecophysiology of gelatinous Nostoc colonies: unprecedented slow growth and survival in resource-poor and harsh environments

Background

The cyanobacterial genus Nostoc includes several species forming centimetre-large gelatinous colonies in nutrient-poor freshwaters and harsh semi-terrestrial environments with extended drought or freezing. These Nostoc species have filaments with normal photosynthetic cells and N₂-fixing heterocysts embedded in an extensive gelatinous matrix of polysaccharides and many other organic substances providing biological and environmental protection. Large colony size imposes constraints on the use of external resources and the gelatinous matrix represents extra costs and reduced growth rates.

Scope

The objective of this review is to evaluate the mechanisms behind the low rates of growth and mortality, protection against environmental hazards and the persistence and longevity of gelatinous Nostoc colonies, and their ability to economize with highly limiting resources.

Conclusions

Simple models predict the decline in uptake of dissolved inorganic carbon (DIC) and a decline in the growth rate of spherical freshwater colonies of N. pruniforme and N. zetterstedtii and sheet-like colonies of N. commune in response to a thicker diffusion boundary layer, lower external DIC concentration and higher organic carbon mass per surface area (CMA) of the colony. Measured growth rates of N. commune and N. pruniforme at high DIC availability comply with general empirical predictions of maximum growth rate (i.e. doubling time 10–14 d) as functions of CMA for marine macroalgae and as functions of tissue thickness for aquatic and terrestrial plants, while extremely low growth rates of N. zetterstedtii (i.e. doubling time 2–3 years) are 10-fold lower than model predictions, either because of very low ambient DIC and/or an extremely costly colony matrix. DIC uptake is limited by diffusion at low concentrations for all species, although they exhibit efficient HCO₃^– uptake, accumulation of respiratory DIC within the colonies and very low CO₂ compensation points. Long light paths and light attenuation by structural substances in large Nostoc colonies cause lower quantum efficiency and assimilation number and higher light compensation points than in unicells and other aquatic macrophytes. Extremely low growth and mortality rates of N. zetterstedtii reflect stress-selected adaptation to nutrient- and DIC-poor temperate lakes, while N. pruniforme exhibits a mixed ruderal- and stress-selected strategy with slow growth and year-long survival prevailing in sub-Arctic lakes and faster growth and shorter longevity in temperate lakes. Nostoc commune and its close relative N. flagelliforme have a mixed stress–disturbance strategy not found among higher plants, with stress selection to limiting water and nutrients and disturbance selection in quiescent dry or frozen stages. Despite profound ecological differences between species, active growth of temperate specimens is mostly restricted to the same temperature range (0–35 °C; maximum at 25 °C). Future studies should aim to unravel the processes behind the extreme persistence and low metabolism of Nostoc species under ambient resource supply on sediment and soil surfaces.     

Elevated CO2 and Phosphate Limitation Favor Micromonas pusilla through Stimulated Growth and Reduced Viral Impact

Growth and viral infection of the marine picoeukaryote Micromonas pusilla was studied under a future-ocean scenario of elevated partial CO₂ (pCO₂; 750 μatm versus the present-day 370 μatm) and simultaneous limitation of phosphorus (P). Independent of the pCO₂ level, the ratios of M. pusilla cellular carbon (C) to nitrogen (N), C:P and N:P, increased with increasing P stress. Furthermore, in the P-limited chemostats at growth rates of 0.32 and 0.97 of the maximum growth rate (μ_max), the supply of elevated pCO₂ led to an additional rise in cellular C:N and C:P ratios, as well as a 1.4-fold increase in M. pusilla abundance. Viral lysis was not affected by pCO₂, but P limitation led to a 150% prolongation of the latent period (6 to 12 h) and an 80% reduction in viral burst sizes (63 viruses per cell) compared to P-replete conditions (4 to 8 h latent period and burst size of 320). Growth at 0.32 μ_max further prolonged the latent period by another 150% (12 to 18 h). Thus, enhanced P stress due to climate change-induced strengthened vertical stratification can be expected to lead to reduced and delayed virus production in picoeukaryotes. This effect is tempered, but likely not counteracted, by the increase in cell abundance under elevated pCO₂. Although the influence of potential P-limitation-relieving factors, such as the uptake of organic P and P utilization during infection, is unclear, our current results suggest that when P limitation prevails in future oceans, picoeukaryotes and grazing will be favored over larger-sized phytoplankton and viral lysis, with increased matter and nutrient flow to higher trophic levels.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Stoichiometry in an ecological context: testing for links between <Emphasis Type="Italic">Daphnia</Emphasis> P-content,growth rate and habitat preference

We used laboratory experiments with ten Daphnia taxa to test for links between Daphnia P-content, growth rate and habitat preference. The taxa represent a wide range of body sizes and most show distinct preferences for one of three habitats: shallow lakes, deep, stratified lakes or fishless ponds. Previous studies show that taxa from shallow lakes and fishless ponds experience high predation risk and rich food resources, whereas taxa from deep lakes experience low predation risk, strong food limitation and potentially P-deficient resources. Thus, we predicted higher P-content and higher maximal growth rates in taxa from ponds and shallow lakes and lower P-content, lower maximal growth but reduced sensitivity to P-limitation in taxa preferring stratified lakes. In each of 25 experiments, a clonal Daphnia cohort was cultured for 4 days on a P-sufficient (molar C:P ratio 70) or a P-deficient (C:P 1,000) diet of a green alga at a high concentration (1 mg C l^–1). The P-content of adult Daphnia fed the P-sufficient diet ranged from 1.52 to 1.22% mass. Small-bodied taxa from shallow lakes had higher P-content than larger-bodied taxa from deep lakes or fishless ponds. However, we found a nonsignificant negative correlation between P-content and growth on the P-sufficient diet, rather than the positive relationship predicted by the growth rate hypothesis. The P-deficient diet resulted in declines in both growth rate and P-content compared with the P-sufficient controls and the extent of the declines differed between taxa. Taxa from ponds showed a marginally greater decline in growth with the P-deficient diet compared with taxa from shallow or deep lakes. However, contrary to stoichiometric theory, no relationship was found between a species P-content and growth depression on the P-deficient diet. Although we found evidence for habitat adaptations, our results show that factors other than Daphnia P-content are important in determining differences between Daphnia species in both maximal growth rate and sensitivity to P-limited growth.     

Effect of Temperature on Growth, Respiration, and Nutrient Regeneration by an Omnivorous Microflagellate

The effect of temperature on the rates and extent of carbon and nitrogen cycling by the heterotrophic microflagellate Paraphysomonas imperforata (diameter, 7 to 12 μm) fed with the diatom Phaeodactylum tricornutum was investigated over an ecologically pertinent temperature range (14 to 26°C). All physiological rates investigated increased with increasing temperature. Q₁₀ values were similar for all rate changes and were comparable to those which have been reported for other protozoa. In contrast to all rates, microflagellate gross growth efficiency and cell volume were unaffected by temperature. Decreases in the concentrations of particulate carbon and particulate nitrogen from grazed diatom cultures also were similar when summed over the entire growth phase of the microflagellate population. Therefore, the proportions of ingested carbon and nitrogen which were incorporated or remineralized by the microflagellate were independent of temperature between 14 and 26°C. At temperatures above 18°C, growth rates of P. imperforata were greater than the maximum growth rates reported for most phytoplankton. We conclude that the impact of P. imperforata on natural phytoplankton communities is not controlled by temperature above 18°C but may be affected by the rate at which zooplankton or microzooplankton prey on the microflagellate, as well as the inability of the microflagellate to graze efficiently when phytoplankton are present at low cell densities.     

Potassium enrichment stimulates the growth and reproduction of a clone of Daphnia dentifera

Nutrient limitation commonly constrains organisms in natural ecosystems. Typically, ecologists focus on limitation by N and P. However, other nutrients can limit growth or reproduction. Here we focus on K limitation of invertebrate consumers (Daphnia dentifera) and phytoplankton in freshwater lakes. All organisms require K for several metabolic processes. In freshwater, K could limit growth because low external concentrations can increase the energetic costs of accumulating K. Furthermore, in a study linking K to disease, we previously found that K enrichment of water from one low-K lake stimulated the growth and reproduction of Daphnia. Here we test whether K could limit the production of Daphnia and phytoplankton across lakes and years. We repeated a life table experiment using water collected from a low-K lake during a different year. K again stimulated Daphnia reproduction. We also enriched water from 12 lakes with K or P and measured short-term growth of Daphnia and the resident algal community. Both nutrients increased Daphnia growth in five lakes. However, only P enhanced algal production. P stimulation of Daphnia positively correlated with algal quantity and the ratio of C to P in seston. However, K stimulation of Daphnia was not correlated with these factors or the background concentration of K. Thus, this study shows repeatable K-limited animal physiology in nature. Further, we can exclude the hypothesis that K stimulates Daphnia indirectly by enhancing algal production. These patterns call for future physiological studies to uncover the mechanistic basis of K limitation in natural systems.     

Increases of Soil C, N, and P Pools Along an Acacia Tree Density Gradient and Their Effects on Trees and Grasses

Nitrogen (N) fixing trees including many species of Acacia are an important though variable component of savanna ecosystems. It is known that these trees enrich the soil with carbon (C) and N, but their effect on the combined C:N:P stoichiometry in soil is less well understood. Theory suggests that they might reduce available phosphorus (P), creating a shift from more N-limited conditions in grass-dominated to more P-limited conditions in tree-dominated sites, which in turn could feed back negatively on the trees’ capacity to fix N. We studied the effects of Acacia zanzibarica tree density upon soil and foliar N:P stoichiometry, and the N₂-fixation rates of trees and leguminous herbs in a humid Tanzanian savanna. Foliar N:P ratios and N₂-fixation rates of trees remained constant across the density gradient, whereas soil C, N and organic P pools increased. In contrast, the N:P ratio of grasses increased and N₂-fixation rates of leguminous herbs decreased with increasing tree density, indicating a shift towards more P-limited conditions for the understory vegetation. These contrasting responses suggest that trees and grasses have access to different sources of N and P, with trees being able to access P from deeper soil layers and perhaps also utilizing organic forms more efficiently.     

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.