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.     

Factors Controlling Phytoplankton Blooms in a Temperate Estuary: Nutrient Limitation and Physical Forcing

A combination of enclosure nutrient enrichment experiments and historical data analysis was used to identify the factors controlling seasonal dynamics and competition of the phytoplankton community in the Curonian lagoon (Southeast Baltic Sea). Experiments using different nutrient (N, P and Si) manipulations were performed in 10-l enclosures for 48 h. Changes in chlorophyll a concentrations, inorganic nutrient concentrations, and plankton cell density were monitored. Results revealed that phytoplankton development in the lagoon is strongly affected by ambient physical factors (wind, temperature). Nutrient limitation, however, also plays an important role in seasonal succession mechanisms showing quite distinct seasonal development patterns. Based on the data, available phytoplankton seasonal succession in the Curonian lagoon could be described as composed by three phases corresponding to different domination and regulatory mechanisms.     

Flux balancing of light and nutrients in a biofilm photobioreactor for maximizing photosynthetic productivity

This article reports a combined experimental and numerical study on the efficient operation of Porous Substrate Bioreactors. A comprehensive model integrating light transport, mass transport, and algal growth kinetics was used to understand the productivity of photosynthetic biofilms in response to delivery rates of photons and nutrients. The reactor under consideration was an evaporation driven Porous Substrate Bioreactor (PSBR) cultivating the cyanobacteria Anabaena variabilis as a biofilm on a porous substrate which delivers water and nutrients to the cells. In an unoptimized experimental case, this reactor was operated with a photosynthetic efficiency of 2.3%, competitive with conventional photobioreactors. Moreover, through a scaling analysis, the location at which the phosphate delivery rate decreased the growth rate to half of its uninhibited value was predicted as a function of microorganism and bioreactor properties. The numerical model along with the flux balancing techniques presented herein can serve as tools for designing and selecting operating parameters of biofilm based cultivation systems for maximum productivity. © 2014 American Institute of Chemical Engineers Biotechnol. Prog., 30:348–359, 2014     

Effects of sediment resuspension on phytoplankton production: teasing apart the influences of light, nutrients and algal entrainment

1. Wind‐induced sediment resuspension can affect planktonic primary productivity by influencing light penetration and nutrient availability, and by contributing meroplankton (algae resuspended from the lake bed) to the water column. We established relationships between sediment resuspension, light and nutrient availability to phytoplankton in a shallow lake on four occasions. 2. The effects of additions of surficial sediments and nutrients on the productivity of phytoplankton communities were measured in 300 mL gas‐tight bottles attached to rotating plankton wheels and exposed to a light gradient, in 24 h incubations at in situ temperatures. 3. While sediment resuspension always increased primary productivity, resuspension released phytoplankton from nutrient limitation in only two of the four experiments because the amount of available nitrogen and phosphorus entrained from the sediments was small compared with typical baseline levels in the water column. In contrast, chlorophyll a entrainment was substantial compared with baseline water column concentrations and the contribution of meroplankton to primary production was important at times, especially when seasonal irradiance in the lake was high. 4. Comparison of the in situ light climate with the threshold of light‐limitation of the phytoplankton indicated that phytoplankton in the lake were only likely to be light‐limited at times of extreme turbidity (e.g. >200 nephelometric turbidity units), particularly when these occur in winter. Therefore, resuspension influenced phytoplankton production mainly via effects on available nutrients and by entraining algae. The importance of each of these varied in time. 5. The partitioning of primary productivity between the water column and sediments in shallow lakes greatly influences the outcome of resuspension events for water column primary productivity.     

Interactions between Light Situation,Depth of Mixing and Phytoplankton Growth during the Spring Period of Full Circulation

Investigations lasting several years at the artificial Lake Saidenbach have shown that, due to the poor light situation expressed in terms of average daily radiation (Riley 1957), mean daily exposure (Reynolds 1973), extinction depth and the ratio ^zmis/^zeu, no mass development of the phytoplankton should actually occur during the spring complete circulation. In practice, however, the period of complete circulation of the water in spring is not only a prerequisite for, but often also the factor causing, the mass development (usually Asterionella formosa). The poor light situation and the fact that the vertical phytoplankton profile shows significant differences between the plankton concentrations at different depths indicate that the spring complete circulation does not represent a complete recirculation, and thus mixing, of the water down to the bottom but involves only episodic and local partial recirculations interspersed with periods of relatively slight turbulence. The actual mixing depth during this period of complete circulation is therefore obviously less than the mean depth of the water concerned, which is commonly assumed to equal. This permits the algae in the upper layers to grow. Respiratory losses of the phytoplankton at greater depths probably remain slight due to their adaptation to low light intensities in winter.     

An analysis,by bioassay,of the factors which limit algal growth in the P. K. le Roux Impoundment,Orange River,South Africa

Factors which limit algal growth in a turbid man-made impoundment were investigated using natural phytoplankton populations. Results indicate that light was the primary limiting factor with nutrients possibly limiting growth in the event of an increase in water transparency. These results differ from another study which used Selenastrum capricornutum bioassays of water from other areas of the Orange River. The value of natural phytoplankton bioassay techniques is supported.     

Phytoplankton diversity loss along a gradient of future warming and brownification in freshwater mesocosms

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Changes in phytoplankton community along a transversal section of the Lower Paraná floodplain,Argentina

Interactions between the main stream of large rivers and their floodplains are usually observed, determining gradual changes in environmental variables along a transversal axis. This paper studies the natural phytoplanktonic community, as well as the main physical and chemical properties from a transversal section of the Lower Paraná River floodplain, encompassing the main channel, a lake and a swamp. In vitro nutrient bioassays for phytoplankton growth were performed in order to evaluate the importance of nutrients as potential limiting factors. A decrease in depth, pH, dissolved oxygen, suspended solids, DIN (dissolved inorganic N), particularly nitrates, and DIN:SRP (soluble reactive P) ratio, was observed from the main channel towards the swamp; as well as an increase in transparency, chlorophyll-a and phytoplankton density. Phytoplankton from the mainstream consisted mostly of typical potamoplanktonic species adapted to grow in turbid environments (mainly Aulacoseira granulata (Müll.) Sim. and its varieties), while in the swamp it was composed mainly of mixotrophic phytoflagellates (euglenoids and cryptomonads), facultative N-heterotrophic diatoms and small-size green algae. Values of both abiotic and biotic variables measured in the lake were always intermediate between the former ones, although closer to those of the river, an account of their permanent connection. Phytoplankton response to N-addition was more frequent in the swamp bioassays than in the other two sites. However, a great increase in algal density in the control flasks, suggests a plentiful source of N at the beginning of the experiments. Light scarcity is more likely to occur than N-limitation in both river and lake due to their low transparency and a high DIN concentration. Although N-limitation may be expected in the swamp, the growth in the controls in spite of the low initial DIN concentration, suggests the existence of an N-supply other than DIN (perhaps organic N). Rather than indicate in situ N-limitation, present results seem, therefore, to suggest that phytoplankton of the swamp mainly composed of mixotrophic algae would be adapted to grow under low DIN concentrations. Possible reasons for the absence of N₂-fixing Cyanobacteria in this environment are discussed.     

Responses of Phytoplankton to in‐situ Nutrient Enrichment; Potential Influences on Species Dominance in a River

The Hawkesbury River at Sackville, New South Wales, Australia is fresh and vigorously mixed by tidal movement. The location has frequent blooms of Microcystis aeruginosa, which have been recorded occurring throughout the year, including winter temperatures as low as 13 °C. Nutrient enrichment tests were performed in‐situ on the natural phytoplankton population in 1997 and 1998 while Microcystis aeruginosa dominated (covering both summer and winter periods). These experiments compared population changes under the ambient nutrient regime with those after additions of ortho‐phosphate, nitrate, ammonia and various combinations of these nutrients. Under ambient conditions, the Microcystis population was able to grow significantly (P < 0.05) while most non‐cyanobacterial phytoplankton did not. Nutrient additions induced a variety of nutrient limitation responses that often varied between genera of major groups i.e. in the Chlorophyceae (Actinastrum sp. responded to phosphorus while Psephonema sp. responded to nitrogen). The possibility that shifts in population dominance from Chlorophyceae to the Cyanobacteria (M. aeruginosa) at Sackville are in response to competition for limiting nutrients is discussed.     

The distribution of soil nutrients with depth: Global patterns and the imprint of plants

To understand the importance of plants in structuring the vertical distributions of soil nutrients, we explored nutrient distributions in the top meter of soil for more than 10,000 profiles across a range of ecological conditions. Hypothesizing that vertical nutrient distributions are dominated by plant cycling relative to leaching, weathering dissolution, and atmospheric deposition, we examined three predictions: (1) that the nutrients that are most limiting for plants would have the shallowest average distributions across ecosystems, (2) that the vertical distribution of a limiting nutrient would be shallower as the nutrient became more scarce, and (3) that along a gradient of soil types with increasing weathering-leaching intensity, limiting nutrients would be relatively more abundant due to preferential cycling by plants. Globally, the ranking of vertical distributions among nutrients was shallowest to deepest in the following order: P > K > Ca > Mg > Na = Cl = SO₄. Nutrients strongly cycled by plants, such as P and K, were more concentrated in the topsoil (upper 20 cm) than were nutrients usually less limiting for plants such as Na and Cl. The topsoil concentrations of all nutrients except Na were higher in the soil profiles where the elements were more scarce. Along a gradient of weathering-leaching intensity (Aridisols to Mollisols to Ultisols), total base saturation decreased but the relative contribution of exchangeable K⁺ to base saturation increased. These patterns are difficult to explain without considering the upward transport of nutrients by plant uptake and cycling. Shallower distributions for P and K, together with negative associations between abundance and topsoil accumulation, support the idea that plant cycling exerts a dominant control on the vertical distribution of the most limiting elements for plants (those required in high amounts in relation to soil supply). Plant characteristics like tissue stoichiometry, biomass cycling rates, above- and belowground allocation, root distributions, and maximum rooting depth may all play an important role in shaping nutrient profiles. Such vertical patterns yield insight into the patterns and processes of nutrient cycling through time.     

Mixotrophy in gyrodinium galatheanum (DINOPHYCEAE): grazing responses to light intensity and inorganic nutrients*

This paper presents results of field and laboratory studies on mixotrophy in the estuarine dinoflagellate Gyrodinium galatheanum (Braarud) Taylor. We tested the hypotheses that this primarily photosynthetic organism becomes phagotrophic when faced with suboptimal light and/or nutrient environments. In Chesapeake Bay, incidence of feeding of this species on cryptophytes is positively correlated with prey density and concentrations of nitrate and nitrite, but negatively correlated with depth, salinity, and phosphate concentration. Feeding in natural assemblages and cultures increased hyperbolically with light intensity. The stoichiometric proportions of dissolved inorganic P and N (DIP:DIN) at the stations where G. galatheanum was present were far below the optimal growth P:N (1:10). Incidence of feeding was negatively related to the ratio of DIP to DIN, suggesting that P limitation may have induced feeding. Addition of nitrate, or addition of both nitrate and phosphate, inhibited feeding in a natural population, indicating that N limitation may also induce feeding. Ingestion of the cryptophyte, Storeatula major, by cultured G. galatheanum was higher in media low in nitrate or phosphate or both, but moderate rates of feeding occurred in nutrient‐replete cultures. When cells were grown in media with varying concentrations of nitrate and phosphate, N deficiency resulted in greater cellular N and Chl a losses than did P deficiency, but P deficiency stimulated feeding more than N deficiency. Both N and P deficiency, or P:N ratios that deviated greatly from 1:10, result in an increase of cellular carbon content and an increase in propensity to feed. Our results suggest that feeding in G. galatheanum is partly a strategy for supplementing major nutrients (N and P) that are needed for photosynthetic carbon assimilation. Feeding in G. galatheanum may also be a strategy for supplementing C metabolism or acquiring trace organic growth factors, since feeding occurs, although at a reduced rate, in nutrient‐replete cultures.     

Phytoplankton assemblages in twenty-one Sicilian reservoirs: relationships between species composition and environmental factors

Data collected in a limnological survey, carried out between 1987 and 1988 on 21 Sicilian reservoirs of varying trophic state, were ordinated using CANOCO 3.1 to generalise the way in which the structure of phytoplankton assemblage is conditioned by both physical and chemical variables. The results showed that in these man-made lakes, characterised by conspicuous water-level fluctuations, the annual and interannual variability in the abundance and composition of phytoplankton may be strongly influenced by their peculiar hydraulic regimes rather than by nutrient availability. In particular, it was highlighted that, from the early summer, water abstraction often leads to increased circulation and to the deepening of the mixed layer. In this way, an increase of the ratio of mixing depth to euphotic depth is forced, with the result that phytoplankton cells experience longer periods in darkness as they are carried through the mixed layer. Phytoplankton assemblages change in species composition in response to the environmental variation. Both the raising of the trophic state, with an increase in phytoplankton biomass and a decrease in transparency, and the intensified abstraction enhance the role of light availability in promoting the development of specific phytoplankton assemblages adapted to the modified physical environment. Light climate is an important influence on the species structure of the phytoplankton, especially in the higher part of the trophic gradient. In contrast, the influence of nutrients on the structure of the assemblages appears to be higher in the lower part of the trophic spectrum or in those environments characterised by a higher hydrological stability during the year.     

Physiological reactions to a change in light regime in cultured Skeletonema costatum (Bacillariophyta): implications for estimation of phytoplankton biomass

The marine planktonic diatom Skeletonema costatum (Cleve) was grown in batch culture under a 12 h light: 12 h dark (LD) regime for several generations before the experiment. At Time 0, half of the culture was transferred to continuous light (CL). Particulate organic carbon (POC), nitrogen (PON), and photosynthetic pigments (measured by spectrophotometry and high performance liquid chromatography) were monitored for 7 to 9 days in the two cultures. Under CL, POC and PON production were always lower. In addition, we measured a much higher proportion of chlorophyll degradation products (as chlorophyllid a, phaeophytin a and phaeophorbid a (PrD)). These indicate different physiological conditions between the two cultures, as reflected also be the lower POC/PON and chlorophyll a/POC ratios. Under CL illumination cells appear stressed, probably due to the total quantity of light to which cells are exposed. In fact, a higher proportion of the two xanthophylls diadinoxanthin and diatoxanthin in the CL culture indicates a photoprotective reaction of the cells. In contrast with the LD culture, parameters measured were not significantly inter-correlated in the CL culture. The lack of correlation between chlorophyll a and POC in CL does not encourage the use of the chlorophyll a/POC ratio as a biomass estimator. On the opposite, the ratio of total pigment content to POC did not vary much as a function of the light condition of the culture, remaining near 0.040 in both light regimes. Although further observations are needed, this index seems to be a reliable indicator of phytoplankton carbon biomass.     

Determining critical light and hydrologic conditions for macrophyte presence in a large shallow lake: The ratio of euphotic depth to water depth

Light determines macrophyte distribution, community composition and biomass in shallow lakes. Therefore, it is vital to determine the critical underwater light climate thresholds for macrophyte degradation and recovery. In this study, we first proposed a novel index, defined as the ratio of euphotic depth (Z_eu) to water depth (WD), as a measure of the underwater light supply for macrophytes. The underwater light environment in Lake Taihu (a large, shallow, eutrophic lake) was then characterized based on this index (Z_eu/WD) using field measurements collected from 2006 to 2013 (8 years × 4 seasons × 32 sites). The distribution of the macrophyte presence frequency (MPF, the number of investigations that identified macrophytes divided by the total number of investigations) was greater than 0.70 in Xukou Bay and East Lake Taihu over the 32 investigations, followed by the other sites distributed in East Lake Taihu. The proportion of macrophyte coverage increased with the increase in Z_eu/WD. A significant relationship was observed between Z_eu/WD and MPF for the 19 sites with macrophytes (r² = 0.48, p < 0.001, n = 19). In the region with high nutrient concentrations and serious water pollution, better underwater light conditions are required for the growth of macrophytes. A Z_eu/WD value of 0.80 can be regarded as the critical underwater light threshold for the growth of macrophytes in Lake Taihu. The region with Z_eu/WD ranging between 0.57 and 0.80 was usually covered by sparse macrophytes; this region should be vital for macrophyte recovery and environmental management in Lake Taihu. The distribution of Z_eu/WD was further obtained using MODIS satellite-derived Z_eu from June to October in 2003 and 2013. Xukou Bay and Guangfu Bay in the southern part of Lake Taihu could be regarded as potentially crucial regions for the recovery of macrophytes from the perspective of underwater light and nutrient levels.     

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.     

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.     

泽雅水库混合深度的年内变化及其对藻类生消影响

就水库藻类水华的发生机制而言,近年来值得关注的研究动向是,一些研究者开始认识到光照和混合的交互作用可能对藻类水华生消过程起到了关键作用,即认为水体混合深度的变化直接影响着可获得光强,从而决定着藻类生物量的时空分布。在这些研究中,混合层深度的确定是探讨藻类生消过程影响的前提。然而已有的研究大都只进行粗略的估计。本文提出采取系统聚类分析的方法确定泽雅水库不同时期的混合深度。在具体分析过程中,将测得的某一天某一深度处的pH,DO,温度,电导率,氧化还原电位等5个参数结果看成一类,采用类间平均连接法,间隔尺度变量为欧氏距离的平方进行聚类。对采用系统聚类分析法和已有的"1°C"法计算得的全年的混合深度的变化趋势进行比较,认为系统聚类分析法可以更为准确地反应水体混合的实际情况。依据混合深度的不同特征,在年内水库水体可划分为热成层时期,全混时期和两者之间的过渡期,藻类生物量与混合深度呈负相关。在热成层期间,藻类生物量都超过了13mm³L^-1。而到了9月份,当混合层深度扩展到15m以上时,藻类生物量明显下降,此后水体处于全混时期,藻类生物量较低,在9-20mm³/L^-1波动。基于稳态假设的前提下,发现随着混合深度增加,稳态藻类生物量呈单峰变化,在混合深度2m处达到最大值,即在混合深度为0-2m之间时,藻类沉降作用是藻类生物量的主要影响因素,随着深度增加,沉降作用下降,因此藻类生物量增加;当混合深度超过2m之后,光限制作用占主导因素,藻类生物量随混合深度增加而下降。这一结果为 Diehl的假说提供了水库现场的实证。     

Photosynthetic responses to dynamic light under field conditions in six tropical rainforest shrubs occuring along a light gradient

 We examined in the field the photosynthetic utilization of fluctuating light by six neotropical rainforest shrubs of the family Rubiaceae. They were growing in three different light environments: forest understory, small gaps, and clearings. Gas exchange techniques were used to analyse photosynthetic induction response, induction maintenance during low-light periods, and lightfleck (simulated sunfleck) use efficiency (LUE). Total daily photon flux density (PFD) reaching the plants during the wet season was 37 times higher in clearings than in the understory, with small gaps exhibiting intermediate values. Sunflecks were more frequent, but shorter and of lower intensity in the understory than in clearings. However, sunflecks contributed one-third of the daily PFD in the understory. Maximum rates of net photosynthesis, carboxylation capacity, electron transport, and maximum stomatal conductance were lower in understory species than in species growing in small gaps or clearings, while the reverse was true for the curvature factor of the light response of photosynthesis. No significant differences were found in the apparent quantum yield. The rise of net photosynthesis during induction after transfer from low to high light varied from a hyperbolic shape to a sigmoidal increase. Rates of photosynthetic induction exhibited a negative exponential relationship with stomatal conductance in the shade prior to the increase in PFD. Leaves of understory species showed the most rapid induction and remained induced longer once transferred to the shade than did leaves of medium- or high-light species. LUE decreased rapidly with increasing lightfleck duration and was affected by the induction state of the leaf. Fully induced leaves exhibited LUEs up to 300% for 1-s lightflecks, while LUE was below 100% for 1–80 s lightflecks in uninduced leaves. Both induced and uninduced leaves of understory species exhibited higher LUE than those of species growing in small gaps or clearings. However, most differences disappeared for lightflecks 10 s long or longer. Thus, understory species, which grew in a highly dynamic light environment, had better capacities for utilization of rapidly fluctuating light than species from habitats with higher light availability. Received: 4 January 1997 / Accepted: 28 April 1997     

Phytoplankton composition and abundance of a central Amazonian floodplain lake

The phytoplankton community of Lake Camaleão, a smallfloodplainlake influenced by a large whitewater river, the Solimões, was monthlyinvestigated for the composition and abundance of itsphytoplankton. The seasonal influence of the floodregime on biomass, species richness and diversity, andits relation with physical and chemical factors(temperature, pH, dissolved oxygen, electricalconductivity, total seston and inorganic nutrients)was analyzed and subjected to principal componentanalysis. Diversity was variable along the seasonalcycle: relatively high values were observed at the endof the dry season supported by high nutrientconcentrations. The phytoplankton was comprised of 262 taxa,with strong dominanceof euglenoids (81%). The three sample stations did not differamong each other, except in the dry season, due todata cluster in relation to theprincipal axis (1 and 2), explaining 63% of thevariation. Biomass accumulation as a function of lakearea reduction contributed to theseresults, indicating that the phytoplankton dynamicswere hydrology-driven.     

Effects of light and microcrustacean prey on growth and investment in carnivory in Utricularia vulgaris

SUMMARY 1. In a 5-week enclosure experiment, we studied the effects of light (ambient light, low light) and prey availability (no prey, prey added) on growth and investment in carnivory in Utricularia vulgaris .
2. Investment in carnivory, measured as the proportion of biomass allocated to bladders, was strongly affected by our manipulations of light intensity and prey density. In the treatment with high prey density the light reduction decreased the investment in bladders from 25% to zero. The effect of prey density on investment in bladders was negative. Because prey addition increased the concentration of nutrients, especially phosphorus, we propose that the effect of the prey treatment on investment reflected altered nutrient concentrations.
3. Availability of prey increased growth and apical biomass of Utricularia . As Utricularia had very few bladders in some treatments we suggest that the effect was due to a combination of live prey trapped and increased nutrient availability from dead prey.
4. Abundance of periphyton on Utricularia and on the enclosure walls was highest in the treatments with high prey density where nutrient concentrations were highest. Thus we interpret the response of periphyton as primarily reflecting nutrient availability.