Nitrogen, macrophytes, shallow lakes and nutrient limitation: resolution of a current controversy?

Geomorphic research across the semi-arid and wet-dry tropics of northern Australia has highlighted the role of the regions’ estuaries as a “canary in the coalmine” for climate variations, with dramatic structural shifts over the Holocene. This behaviour highlights the region’s potential sensitivity to climate change and suggests the need for careful identification and interpretation of dynamics in the tropical and sub-tropical regions. However, analysis of change in these regions requires care, as a number of the basic tools applied to interpreting estuarine change in temperate regions are obscured, invalid or simply unavailable when applied in lower latitudes. This study provides a synthesis from a range of investigations across northern Australia and identifies characteristics to be considered when interpreting or predicting sub-millennial estuarine change in these regions.     

Seasonal dynamics of chemotypes in a freshwater phytoplankton community – A metabolomic approach

Cyanobacteria are known to produce a huge variety of secondary metabolites. Many of these metabolites are toxic to zooplankton, fish, birds and mammals. Therefore, the toxicity of cyanobacterial blooms is strongly dependent on the cyanobacterial strain composition. These strains produce distinct bouquets of secondary metabolites, and thus constitute different chemotypes. Some of the cyanobacterial metabolites are potent inhibitors of gut proteases of the filter-feeder Daphnia. Here, we investigate the seasonal dynamics of secondary metabolites in a phytoplankton community from a hypertrophic pond, making use of a new metabolomic approach. Using liquid chromatography coupled with high-resolution mass spectrometry (LCMS), we obtained mass spectra of phytoplankton samples taken on different dates throughout the summer season. By applying multivariate statistics, we combined these data with the protease inhibition capacity of the same samples. This led to the identification of metabolites with cyanobacterial origin and as well of distinct cyanobacterial chemotypes being dominant on different dates. The protease inhibition capacity varied strongly with season, and only one out of 73 known cyanobacterial protease inhibitors could be confirmed in the natural samples. Instead, several so far unknown, putative protease inhibitors were detected. In conclusion, the creation of time series of mass spectral data of a natural phytoplankton community proved to be useful for elucidating seasonal chemotype succession in a cyanobacterial community. Additionally, correlating mass spectral data with a biological assay provides a promising tool for facilitating the search for new harmful metabolites prior to structure elucidation.     

Rehabilitation of shallow lakes: time to adjust expectations?

We investigated the relative importance of flood defenses and other environmental variables for the cladoceran and copepod communities in floodplain water bodies in Southeastern Norway. The water bodies covered gradients of size, distance to the river and water chemistry, and half of them were located behind flood defenses. The effects of environmental variables on the communities were analyzed using redundancy analysis. The set of environmental variables accounted for more of the explained variation in the cladoceran community than in the copepod community. Water quality was much more important than flooding-related variables for both communities. Although cladoceran species richness was slightly higher in water bodies outside flood defenses, total nitrogen, total organic carbon (TOC), and water body area were the most important factors for the cladocerans. Macrophyte coverage was the most important variable for both species richness and community structure of copepods. Although our results show that water quality and spatial/structural variables are more important than flood defenses in structuring the communities, the effect of TOC on the cladoceran community could likely be mediated through effects of flood defense on TOC. Prospects for a continued anthropogenic pressure on river floodplains raise concern for the future of these unique ecosystems.     

Response of a shallow Mediterranean lake to nutrient diversion: does it follow similar patterns as in northern shallow lakes?

1. In view of the paucity of data on the response of warm shallow lakes to reductions in nutrient loading, this paper presents a long‐term limnological data set to document changes in the food‐web of a shallow Mediterranean lake (Lake Albufera, Valencia, Spain) that has experienced reductions in phosphorus (P) (77%) and nitrogen (N) (24%) loading following sewage diversion. 2. Nine years after sewage diversion, P concentration in the lake was reduced by 30% but remained high (TP = 0.34 mg L^?1), although the mean water retention time in the lake was only 0.1 years. Nitrate concentrations did not significantly change, probably because the lake continued to receive untreated effluents from ricefields. 3. Chlorophyll a concentration was reduced by half (annual mean of 180 μg L^?1). Cyanobacteria abundance remained high but its composition changed towards smaller species, both filamentous and chroococcal forms. 4. Cladocera abundance increased and reached peaks twice a year (December to March and July to September). After nutrient reduction, short‐term clear‐water phases (up to 5 weeks) occurred during February to March in several years, concomitant with annual flushing of the lake and lower fish densities. The abundance of Cladocera in winter contrasted with the spring peaks observed in northern restored shallow lakes. The zooplankton to phytoplankton biomass ratio remained lower than in northern temperate shallow lakes, probably because of fish predation on zooplankton. 5. Improvement of the water quality of Lake Albufera remained insufficient to counteract littoral reed regression or improve underwater light allowing submerged plants re‐colonise the lake. 6. Sewage diversion from Lake Albufera impacted the food web through the plankton, but higher trophic levels, such as fish and waterfowl, were affected to a lesser degree. Although the fish species present in the lake are mainly omnivorous, long‐term data on commercial fish captures indicated that fish communities changed in response to nutrient level and trophic structure as has been observed in restored shallow lakes at northern latitudes. 7. Phosphorus concentrations produced similar phytoplankton biomass in Lake Albufera as in more northern shallow lakes with abundant planktivorous fish and small zooplankton. However, in Lake Albufera, high average concentrations were maintained throughout the year. Overall, results suggest that nutrient control may be a greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes.     

Is roach herbivory preventing re-colonization of submerged macrophytes in a shallow lake?

Hydrobiologia - Re-establishing a stable submerged vegetation is considered an important tool to restore shallow eutrophic lakes. Enhanced turbidity and sediment re-suspension as well as grazing by...     

Triplet state dynamics in peridinin-chlorophyll-a-protein: a new pathway of photoprotection in LHCs?

This work investigates the interaction of carotenoid and chlorophyll triplet states in the peridinin-chlorophyll-a-protein (PCP) of Amphidinium carterae using step-scan Fourier transform infrared spectroscopy. We identify two carotenoid triplet state lifetimes of approximately 13 and approximately 42 mus in the spectral region between 1800 and 1100 cm(-1) after excitation of the 'blue' and 'red' peridinin (Per) conformers and the Q(y) of chlorophyll-a (Chl-a). The fast and slow decaying triplets exhibit different spectral signatures in the carbonyl region. The fast component generated at all excitation wavelengths is from a major conformer with a lactone stretching mode bleach at 1745 cm(-1). One (1720 cm(-1)) and two (1720 cm(-1) and 1741 cm(-1)) different Per conformers are observed for the slow component upon 670- and 530-480-nm excitation, respectively. The above result implies that (3)Per triplets are formed via two different pathways, corroborating and complementing visible triplet-singlet (T-S) spectra (Kleima et al., Biochemistry (2000), 39, 5184). Surprisingly, all difference spectra show that Per and Chl-a modes are simultaneously present during the (3)Per decay, implying significant involvement of (3)Chl-a in the (3)Per state. We suggest that this Per-Chl-a interaction via a delocalized triplet state lowers the (3)Per energy and thus provides a general, photoprotection mechanism for light-harvesting antenna complexes.     

Spatial structure and β-diversity of phytoplankton in Tibetan Plateau lakes: nestedness or replacement?

Hydrobiologia - Spatial patterns and β-diversity of phytoplankton assemblages depend on the relative importance of species dispersal capacity and species-sorting. Variability in species...     

The contribution of size-fractionated plankton to biomass and primary production of phytoplankton in saline–alkaline ponds

Primary productivity, biomass and chlorophyll-a of size fractionated phytoplankton (<0.22 m, <3 m, <8 m, <10 m, <40 m, <64 m, <112 m and <200 m) were estimated in 6 ponds and 5 experimental enclosures. The results showed that the planktonic algae less than 10 m are important in the biomass and production of phytoplankton in saline–alkaline ponds. The production of size fractionated phytoplankton corresponding to <112 m, <10 m and <3 m in saline–alkaline ponds were 10.5 ± 6.6 , 8.6 ± 5.4 and 0.33 ± 0.1 mgC l^–1 d^–1, respectively. Mean community respiration rate was 1.80 ± 0.73, 1.69 ± 0.90 and 1.38 ± 1.12 mgC l^–1 d^–1, respectively. The average production of phytoplankton corresponding to micro- (10–112 m), nano- (3–10 m) and pico- (<3 m) were 1.61, 8.30 and 0.33 mgC l^–1 d^–1, respectively. The ratio of those to the total phytoplankton production was 15%, 79% and 3%, respectively. The mean respiration rate of the different size groups was 0.11, 0.31 and 1.38 mgC l^–1 d^–1; the ratio of those to total respiration of phytoplankton was 6%, 17% and 77%, respectively. The production of size-fractionated phytoplankton corresponding to <200 m, <10 m and <3 m in enclosures was 2.19 ± 1.63, 2.08 ± 1.75 and 0.22 ± 0.08 mgC l^–1 d^-1, respectively. Mean community respiration rates were 1.25 ± 1.55, 1.17 ± 1.42 and 0.47 ± 0.32 mgC l^–1 d^–1, respectively. The average production of phytoplankton corresponding to micro- (10–200 m), nano- (3–10 m) and pico- (<3 m) plankton was 0.11, 1.86 and 0.22 mgC l^–1 d^–1, respectively. The ratio of those to the total production of phytoplankton was 5%, 85% and 10%, respectively. The mean respiration rate of different size groups were 0.08, 0.72 and 0.46 mgC l^–1 d^–1, the ratio of those to total respiration of phytoplankton was 6%, 57% and 37%, respectively. The concentrations of chlorophyll-a of the phytoplankton in the corresponding size of micro- (10–112 m), nano- (3–10 m) and pico- (<3 m) plankton in the experimental ponds were 19.3, 98.2 and 11. 9 g l^–1, respectively. The ratio of those to the total chlorophyll-a was 15%, 76% and 9%, respectively. The concentrations of chlorophyll-a of phytoplankton micro- (10–200 m), nano- (3–10 m) and pico- (<3 m) plankton in enclosures were 1.7, 34.3 and 3.0 g l^–1, respectively. The ratio of those to the total chlorophyll-a was 4%, 88% and 8%, respectively.     

Primary production of aquatic macrophytes and their epiphytes in two shallow lakes (Peipsi and Võrtsjärv) in Estonia

In shallow lakes with large littoral zones, epiphytes and submerged macrophytes can make an important contribution to the total annual primary production. We investigated the primary production (PP) of phytoplankton, submerged macrophytes, and their epiphytes, from June to August 2005, in two large shallow lakes. The production of pelagic and littoral phytoplankton and of the dominant submerged macrophytes in the littoral zone (Potamogeton perfoliatus in Lake Peipsi and P. perfoliatus and Myriopyllum spicatum in Lake Võrtsjärv) and of their epiphytes was measured using a modified ¹⁴C method. The total PP of the submerged macrophyte area was similar in both lakes: 12.4 g C m^?2 day^?1 in Peipsi and 12.0 g C m^?2 day^?1 in Võrtsjärv. In Peipsi, 84.2% of this production was accounted for by macrophytes, while the shares of phytoplankton and epiphytes were low (15.6 and 0.16%, respectively). In Võrtsjärv, macrophytes contributed 58%, phytoplankton 41.9% and epiphytes 0.1% of the PP in the submerged macrophyte area. Epiphyte production in both lakes was very low in comparison with that of phytoplankton and macrophytes: 0.01, 5.04, and 6.97 g C m^?2 day^?1, respectively, in Võrtsjärv, and 0.02, 1.93, and 10.5 g C m^?2 day^?1, respectively, in Peipsi. The PP of the littoral area contributed 10% of the total summer PP of Lake Peipsi sensu stricto and 35.5% of the total summer PP of Lake Võrtsjärv.     

Seasonal changes in cell‐mediated immunocompetence and mass gain in nestling barn swallows: a parasite‐mediated effect?

Reproduction in seasonal environments is usually timed so peak demand for food by offspring coincides with peak availability. Hence, late breeders will encounter a scarcity of food. Since parasite populations grow during the reproductive season of their hosts, late reproducing animals will also face an increasing challenge by parasites. We hypothesised that seasonal decrease in food availability and seasonal increase in parasite abundance will cause a trade-off between growth and immune function. This prediction was tested in nestling barn swallows ( Hirundo rustica ) from first and second broods. Nestlings from second broods mounted stronger T cell mediated immune responses to a challenge with a novel antigen, but had lower rates of mass gain, than nestlings from first broods, consistent with the prediction. Broods in which at least one nestling died had lower levels of T cell mediated immune response, but not lower rates of mass gain, than broods without mortality, suggesting that brood reduction is mediated through an inability of offspring to defend themselves against parasites rather than an inability to grow. Possible mechanisms include scarcity of specific nutrients needed for immune responses, and/or parasites being concentrated on a single or few nestlings.     

How important is the crustacean plankton for the maintenance of water clarity in shallow lakes with abundant submerged vegetation?

• 1 We measured the abundance and biomass of filter‐feeding microcrustacean zooplankton and calculated their grazing impact on phytoplankton biomass during summer in five shallow, mesotrophic to eutrophic lakes. For three of the lakes data exist both from years with dense submerged vegetation and low turbidity (the clearwater state), as well as from years characterised by sparse vegetation and high turbidity (the turbid state). In the other two lakes data are available only for clearwater conditions.
• 2 In all lakes conditions of dense vegetation and clear water coincided with a low abundance of crustacean plankton during summer. In the three lakes that shifted, the calculated biovolume ingested by crustacean plankton (filtering rate) was 3–11 times lower during clearwater conditions compared with turbid conditions. Because phytoplankton biomass was lower during clearwater conditions, however, daily grazing pressure from microcrustacea (expressed as percentage of phytoplankton biomass) did not differ between states. In three of the five lakes, grazers were estimated to take less than 10% of the phytoplankton biomass per day, indicating filtration by zooplankton was not the most important mechanism to maintain clearwater conditions.
• 3 High densities of Cladocera were found in three of the lakes within dense stands of Charophyta. However, these samples were dominated by plant‐associated taxa that even during the night were rarely found outside the vegetation. This indicates that plant‐associated zooplankton has no major influence on the maintenance of water clarity outside the vegetation.
• 4 Spring peak abundance of Cladocera was observed in three of the lakes. In two of these, where seasonal development was studied in both the clearwater and the turbid state, spring peaks were lower during the clearwater state.
• 5 Predation, low food availability or a combination of both may explain the low zooplankton densities. Phytoplankton may be limited by low phosphorus availability in the lakes dominated by Charophyta. Our results indicate that the importance of zooplankton grazing may have minor importance for the maintenance of the clearwater state in lakes with dense, well‐established submerged vegetation.

     

Seasonal variation in phytoplankton composition and physical-chemical features of the shallow Lake Doïrani,Macedonia, Greece

Phytoplankton species composition, seasonal dynamics and spatial distribution in the shallow Lake Doïrani were studied during the growth season of 1996 along with key physical and chemical variables of the water. Weak thermal stratification developed in the lake during the warm period of 1996. The low N:P ratio suggests that nitrogen was the potential limiting nutrient of phytoplankton in the lake. In the phytoplankton of the lake, Chlorophyceae were the most species-rich group followed by Cyanophyceae. The monthly fluctuations of the total phytoplankton biomass presented high levels of summer algal biomass resembling that of other eutrophic lakes. Dinophyceae was the group most represented in the phytoplankton followed by Cyanophyceae. Diatomophyceae dominated in spring and autumn. Nanoplankton comprised around 90% of the total biomass in early spring and less than 10% in summer. The seasonal dynamics of phytoplankton generally followed the typical pattern outlined for other eutrophic lakes. R-species (small diatoms), dominant in the early phase of succession, were replaced by S-species (Microcystis, Anabaena, Ceratium) in summer. With cooling of the water in September, the biomass of diatoms (R-species) increased. The summer algal maxima consisted of a combination of H and M species associations (sensu Reynolds). Phytoplankton development in 1996 was subject to the combined effect of the thermal regime, the small depth of mixing and the increased sediment-water interactions in the lake, which caused changes in the underwater light conditions and nutrient concentrations.     

Is the island biogeography model a poor predictor of biodiversity patterns in shallow lakes?

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The evolutionary significance of parasitism: do parasite‐driven genetic dynamics occur ex silico?

It has long been recognized that reciprocal antagonism might lock host and parasite populations into a process of constant change, adapting and reacting in open‐ended coevolution. A significant body of theory supports this intuition: dynamic genetic polymorphisms are a common outcome of computer simulations of host–parasite coevolution. These in silico experiments have also shown that dynamical interactions could be responsible for high levels of genetic diversity in host populations, and even be the principle determinant of rates of genetic recombination and sexuality. The evolutionary significance of parasitism depends on the strength and prevalence of parasite‐mediated selection in nature. Here I appraise whether parasitism is a pervasive agent of evolutionary change by detailing empirical evidence for selection. Although there is considerable evidence of genetic variation for resistance, and hence the potential for selection, direct observation of parasite‐driven genetic change is lacking.     

Phenolic content and growth of wetland macrophytes: Is the allocation to secondary compounds driven by nutrient availability?

This study compares soluble phenolics and lignin content in two wetland macrophytes with contrasting life strategies grown under a varying nutrient supply in the field and in a greenhouse experiment. The differences are explained in terms of the protein competition model (PCM) hypothesis relating changes in secondary metabolites to changing nutrient limitation. The two study species, Eleocharis cellulosa (EC) and Typha domingensis (TD), are both widespread in tropical and subtropical freshwater and brackish marshes of the New World, and are often found in P-limited rather than N-limited conditions. TD is a fast-growing competitor with large nutrient requirements. EC is a stress tolerator, quite well adapted to growth in nutrient-limiting environments. In both species, the concentration of phenolics was negatively correlated with increasing growth (due to increasing nutrient levels). This is in agreement with the PCM hypothesis, which predicts an increase in phenolic synthesis when protein synthesis (and consequently growth) is low due to limited resource availability. An interesting difference was found in the correlation between tissue nutrients and phenolics. TD from both the field and the greenhouse showed a negative correlation between tissue P and phenolics, while EC displayed a significant negative correlation between tissue N and phenolics. EC is adapted to low P, and increased tissue P content represents luxury consumption (uptake of P for storage) which is not reflected in increased growth and thus is not correlated with phenolics. These are the first steps in elucidating the relationship among nutrient availability, growth and phenolic content in two important primary producers of tropical and subtropical marshes.     

Correlated evolution of fruit and leaf size in bird‐dispersed plants: species‐level variance in fruit traits explained a bit further?

The astounding morphological diversity exhibited by the fruits of vertebrate-dispersed plants has been traditionally interpreted as the adaptive outcome of divergent selective pressures exerted on plants by the broad array of frugivorous animals involved in seed dispersal. Although the selective capacity of frugivores provides support to this interpretation, recent studies have challenged it by documenting a strong phylogenetic component associated to interspecific variation in most fruit characteristics. Size-related fruit traits provide a conspicuous exception to this pattern, because they exhibit considerable variation at the between-species level which is largely independent of phylogeny and is correlated with consumption by differently-sized dispersal agents. Substantial species-level variance in size-related traits may reflect genuine disperser-driven diversification, but may also be partly influenced by correlated evolution of fruit size with the size of other plant structures. This latter possibility is tested here for bird-dispersed plants of the Iberian Peninsula using phylogenetically independent contrasts. Results demonstrate the existence of correlated evolution of fruit and leaf size at the species level. As all the plant taxa considered have their fruits eaten, and seeds dispersed, by the same relatively reduced set of frugivorous bird species, results suggest that a significant fraction of the variation in fruit size represented in the species sample may be explained as an indirect consequence of variation in leaf size, rather than being associated with adaptive divergence related to seed dispersal agents.     

Species‐specific stomatal response of trees to drought – a link to vegetation dynamics?

Question: Is stomatal regulation specific for climate and tree species, and does it reveal species‐specific responses to drought? Is there a link to vegetation dynamics? Location: Dry inner alpine valley, Switzerland Methods: Stomatal aperture (θ_E) of Pinus sylvestris, Quercus pubescens, Juniperus communis and Picea abies were continuously estimated by the ratio of measured branch sap flow rates to potential transpiration rates (adapted Penman‐Monteith single leaf approach) at 10‐min intervals over four seasons. Results: θ_E proved to be specific for climate and species and revealed distinctly different drought responses: Pinus stomata close disproportionately more than neighbouring species under dry conditions, but has a higher θ_E than the other species when weather was relatively wet and cool. Quercus keeps stomata more open under drought stress but has a lower θ_E under humid conditions. Juniperus was most drought‐tolerant, whereas Picea stomata close almost completely during summer. Conclusions: The distinct microclimatic preferences of the four tree species in terms of θ_E strongly suggest that climate (change) is altering tree physiological performances and thus species‐specific competitiveness. Picea and Pinus currently live at the physiological limit of their ability to withstand increasing temperature and drought intensities at the sites investigated, whereas Quercus and Juniperus perform distinctly better. This corresponds, at least partially, with regional vegetation dynamics: Pinus has strongly declined, whereas Quercus has significantly increased in abundance in the past 30 years. We conclude that θ_E provides an indication of a species' ability to cope with current and predicted climate.     

Interactions between phytoplankton,zooplankton and fish in a shallow,hypertrophic lake: a study of phytoplankton collapses in Lake Søbygård,Denmark

Since 1983 severe phytoplankton collapses have occurred 1–4 times every summer in the shallow and hypertrophic Lake Søbygård, which is recovering after a ten-fold decrease of the external phosphorus loading in 1982. In July 1985, for example, chlorophyll a changed from 650 µg l^–1 to about 12 µg 1^–1 within 3–5 days. Simultaneously, oxygen concentration dropped from 20–25 mg O₂l^–1 to less than 1 mg O₂l^–1, and pH decreased from 10.7 to 8.9. Less than 10 days later the phytoplankton biomass had fully recovered. During all phytoplankton collapses the density of filter-feeding zooplankton increased markedly, and a clear-water period followed. Due to marked changes in age structure of the fish stock, different zooplankton species were responsible for the density increase in different years, and consequently different collapse patterns and frequencies were observed.The sudden increase in density of filter-feeding zooplankton from a generally low summer level to extremely high levels during algae collapses, which occurred three times from July 1984 to June 1986, could neither be explained by changes in regulation from below (food) nor from above (predation). The density increase was found after a period with high N/P ratios in phytoplankton or nitrate depletion in the lake. During that period phytoplankton biomass, primary production and thus pH decreased, the latter from 10.8–11.0 to 10.5. We hypothesize that direct or indirect effects of high pH are important in controlling the filter-feeding zooplankton in this hypertrophic lake. Secondarily, this situation affects the trophic interactions in the lake water and the net internal loading of nutrients. Consequently, not only a high content of planktivorous fish but also a high pH may promote uncoupling of the grazing food-web in highly eutrophic shallow lakes, and thereby enhance eutrophication.A tentative model is presented for the occurrence of collapses, and their pattern in hypertrophic lakes with various fish densities.