Foraging of multimammate mice, Mastomys natalensis, under different predation pressure: cover, patch-dependent decisions and density-dependent GUDs

Patch use under predation risk often results in a change of feeding behaviour in the prey animals. However, such changes only appear if the animals are able to assess under which predation pressure they live. We investigated patch use of Mastomys natalensis under different conditions of avian predation pressure.
In replicated maize field plots in Morogoro, Tanzania, avian predators were allowed under natural conditions (control), attracted with perches and nest boxes or kept out with nets. During four one‐week periods in late 1999, we measured rodent feeding decisions with the giving‐up density (GUD) method. Trays with known amounts of millet seeds in sand were placed in pairs, one of them under a cover, the other one in the open. M. natalensis mice were expected to give up sooner in the open trays than in those with cover. We hypothesised that M. natalensis mice could assess the ambient predation pressure leading to larger difference in GUD between covered and non‐covered trays in the plots where predators were attracted. We also made video recordings of the rodent activity at a pair of trays in each treatment. The GUD‐values were significantly lower for the covered trays but predation pressure did not affect this difference. The video observations showed that in the control and netted plots the animals visited trays equally frequently regardless of the cover, while the visits in the predator‐attracted plots occurred significantly more often in the covered trays. We conclude that M. natalensis can assess the ambient predation pressure and adapt its behaviour at a feeding patch. However, the variation in predation pressure in our experiment was not obvious from the GUD. Moreover, we found a strong relation between rodent density and GUD, which may mask variations in perceived predation pressure. Similar GUD values may be reached in different ways and we present models to investigate whether animals’ decision to forage at a food patch is only affected by the seed density at that patch, not by that at a neighbour patch.     

Reaeration of Oxygen in Shallow,Macrophyte Rich Streams: I – Determination of the Reaeration Rate Coefficient

The rate coefficient K₂ for the exchange of oxygen between flowing water and the atmosphere (reaeration) has been studied in six Danish streams covering a relatively wide range of hydraulic conditions, pollutional loading, and macrophyte abundance. 103 K₂-measurements were performed in 1978–85. 82 measurements were obtained applying 5 different indirect methods all balancing the sources and sinks of stream dissolved oxygen under conditions of normal operation of the system (3 methods) and under artificial depletion of the oxygen concentration of the stream water by addition of sodium sulphite (2 methods). 21 K₂-values were determined directly applying a gaseous tracer (krypton-85) for reaeration. Guidelines for selecting a proper method to determine K₂ knowing macrophyte biomass and loading characteristics of the particular stream are provided.     

Fish manipulation as a lake restoration tool in shallow,eutrophic, temperate lakes 2: threshold levels,long-term stability and conclusions

In order to evaluate short-term and long-term effects of fish manipulation in shallow, eutrophic lakes, empirical studies on relationships between lake water concentration of total phosphorus (P) and the occurrence of phytoplankton, submerged macrophytes and fish in Danish lakes are combined with results from three whole-lake fish manipulation experiments. After removal of less than 80 per cent of the planktivorous fish stock a short-term trophic cascade was obtained in the nutrient regimes, where large cyanobacteria were not strongly dominant and persistent. In shallow Danish lakes cyanobacteria were the most often dominating phytoplankton class in the P-range between 200 and 1 000μg P l⁻¹. Long-term effects are suggested to be closely related to the ability of the lake to establish a permanent and wide distribution of submerged macrophytes and to create self-perpetuating increases in the ratio of piscivorous to planktivorous fish. The maximum depth at which submerged macrophytes occurred, decreased exponentially with increasing P concentration. Submerged macrophytes were absent in lakes>10 ha and with P levels above 250–300μg P l⁻¹, but still abundant in some lakes<3 ha at 650μg P l⁻¹. Lakes with high cover of submerged macrophytes showed higher transparencies than lakes with low cover aboveca. 50μg P l⁻¹. These results support the alternative stable state hypothesis (clear or turbid water stages). Planktivorous fish>10 cm numerically contributed more than 80 per cent of the total planktivorous and piscivorous fish (>10 cm) in the pelagical of lakes with concentrations above 100μg P l⁻¹. Below this threshold level the proportion of planktivores decreased markedly toca. 50 per cent at 22μg P l⁻¹. The extent of the shift in depth colonization of submerged macrophytes and fish stock composition in the three whole-lake fish manipulations follows closely the predictions from the relationships derived from the empirical study. We conclude that a long-term effect of a reduction in the density of planktivorous fish can be expected only when the external phosphorus loading is reduced to below 0.5–2.0 g m⁻² y⁻¹. This loading is equivalent to an in-lake summer concentration below 80–150μg P l⁻¹. Furthermore, fish manipulation as a restoration tool seems most efficient in shallow lakes.     

Differential structural properties of GLP-1 and exendin-4 determine their relative affinity for the GLP-1 receptor N-terminal extracellular domain

Glucagon-like peptide-1 (GLP-1) and exendin-4 (Ex4) are homologous peptides with established potential for treatment of type 2 diabetes. They bind and activate the pancreatic GLP-1 receptor (GLP-1R) with similar affinity and potency and thereby promote insulin secretion in a glucose-dependent manner. GLP-1R belongs to family B of the seven transmembrane G-protein coupled receptors. The N-terminal extracellular domain (nGLP-1R) is a ligand binding domain with differential affinity for Ex4 and GLP-1: low affinity for GLP-1 and high affinity for exendin-4. The superior affinity of nGLP-1R for Ex4 was previously explained by an additional interaction between nGLP-1R and the C-terminal Trp-cage of Ex4. In this study we have combined biophysical and pharmacological approaches thus relating structural properties of the ligands in solution to their relative binding affinity for nGLP-1R. We used both a tracer competition assay and ligand-induced thermal stabilization of nGLP-1R to measure the relative affinity of full length, truncated, and chimeric ligands for soluble refolded nGLP-1R. The ligands in solution and the conformational consequences of ligand binding to nGLP-1R were characterized by circular dichroism and fluorescence spectroscopy. We found a correlation between the helical content of the free ligands and their relative binding affinity for nGLP-1R, supporting the hypothesis that the ligands are helical at least in the segment that binds to nGLP-1R. The Trp-cage of Ex4 was not necessary to maintain a superior helicity of Ex4 compared to GLP-1. The results suggest that the differential affinity of nGLP-1R is explained almost entirely by divergent residues in the central part of the ligands: Leu10-Gly30 of Ex4 and Val16-Arg36 of GLP-1. In view of our results it appears that the Trp-cage plays only a minor role for the interaction between Ex4 and nGLP-1R and for the differential affinity of nGLP-1R for GLP-1 and Ex4.     

Salinity Induced Regime Shift in Shallow Brackish Lagoons

In brackish lagoons, Daphnia is replaced by calanoid copepods (Eurytemora affinis, Acartia spp.) and rotifers when a certain threshold (depending on, for instance, fish density) is reached. We hypothesize that loss of Daphnia induces a regime shift from clear to turbid at high nutrient concentrations. We conducted a factorial designed enclosure experiment with contrasting salinities (0–16‰), low fish predation (one three-spined stickleback, Gasterosteus aculeatus, m⁻²) and three levels of nutrient loading in a shallow brackish lagoon. A change point analysis suggests a strong regime shift from a clear to a turbid state at 6–8‰ salinity at low and high loading, but not for the control. From the low to the high salt regime, chlorophyll a (Chla), Chla:total phosphorus (TP) and Chla:total nitrogen (TN) ratios shifted highly significantly for all nutrient treatments, and the bacterioplankton production followed the changes in Chla. These changes occurred parallel with a shift from cladoceran and cyclopoid copepod to rotifer dominance. Monitoring data from 60 Danish brackish lagoons show increasing Chla with increasing TP and TN as well as interactive effects of TN and salinity, peaking at intermediate salinity. A relatively weak effect of salinity at low nutrient concentrations and the stronger effect at intermediate high salinity are in accordance with the experimental results. However, these data suggest a lower salinity threshold than in the experiment, which may be explained by a higher fish density. Our results have implications for the management of coastal lagoons both at present and in a future (predicted) warmer climate: (1) improved water quality can be obtained by reducing the nutrient loading or enhancing the freshwater input to a level triggering a shift to Daphnia dominance (typically <2‰), (2) fish manipulation is probably not a useful tool for brackish lagoons, unless the salinity is below the threshold for a potential shift to a clear Daphnia dominated state, and (3) more abrupt changes will expectedly occur in low-saline coastal lagoons at increasing salinity during summer in a future warmer climate.     

Impacts of climate warming on lake fish community structure and potential effects on ecosystem function

Fish play a key role in the trophic dynamics of lakes, not least in shallow systems. With climate warming, complex changes in fish community structure may be expected owing to the direct and indirect effects of temperature, and indirect effects of eutrophication, water-level changes and salinisation on fish metabolism, biotic interactions and geographical distribution. We review published and new data supporting the hypotheses that, with a warming climate, there will be changes in: fish community structure (e.g. higher or lower richness depending on local conditions); life history traits (e.g. smaller body size, shorter life span, earlier and less synchronised reproduction); feeding mode (i.e. increased omnivory and herbivory); behaviour (i.e. stronger association with littoral areas and a greater proportion of benthivores); and winter survival. All these changes imply higher predation on zooplankton and macroinvertebrates with increasing temperatures, suggesting that the changes in the fish communities partly resemble, and may intensify, the effects triggered by eutrophication. Modulating factors identified in cold and temperate systems, such as the presence of submerged plants and winter ice cover, seem to be weaker or non-existent in warm(ing) lakes. Consequently, in the future lower nutrient thresholds may be needed to obtain clear-water conditions and good ecological status in the future in currently cold or temperate lakes. Although examples are still scarce and more research is needed, we foresee biomanipulation to be a less successful restoration tool in warm(ing) lakes without a strong reduction of the nutrient load.     

Persistent internal phosphorus loading during summer in shallow eutrophic lakes

Nutrient availability, in particular of phosphorus (P), is a key factor for the structure and functioning of shallow lakes, and not least the sediment plays an important role by acting as both a nutrient source and sink. We used 21 years of monthly mass balance and lake water data from six shallow (mean depth = 1.2–2.7 m) and fast flushed (mean hydraulic retention time = 0.6–2.6 months) eutrophic Danish lakes (mean summer P concentrations ranging from 0.09 to 0.61 mg/l) to investigate long-term trends in yearly and seasonal patterns of P retention. To one of the lakes, the external P input was reduced by 70% in the early 1990s, whereas none of the other lakes have experienced major changes in external P loading for more than 20 years. All lakes showed a distinct seasonal pattern with high P concentrations and typically negative P retention during summer (up to ?300% of the external loading from May to August). During winter, P retention was overall positive (up to 50% of the external loading from December to April). Internal P loading from the sediment delayed lake recovery by approximately 10 years in the lake with the most recently reduced external loading, but in all the lakes net release of P from the sediment occurred during summer. P release in the six lakes has not abated during the past decade, indicating that the sediment of eutrophic and turbid shallow lakes remains a net source of P during summer. The seasonal variations in P retention became more pronounced with increasing P levels, and retention decreased with increasing temperature, but increased if clear water conditions were established.     

Winter ecology of shallow lakes: strongest effect of fish on water clarity at high nutrient levels

While the structuring role of fish in lakes is well studied for the summer season in North temperate lakes, little is known about their role in winter when fish activity and light irradiance potentially are lower. This is unfortunate as the progressing climate change may have strong effects on lake winter temperature and possibly on trophic dynamics too. We conducted an enclosure experiment with and without the presence of fish throughout winter in two shallow lakes with contrasting phosphorus concentrations. In hypertrophic Lake Søbygård, absence of fish led to higher biomass of zooplankton, higher grazing potential (zooplankton:phytoplankton ratio) and, accordingly, lower biomass of phytoplankton and chlorophyll a (Chl a), while the concentrations of total nitrogen (TN), total phosphorus (TP), oxygen and pH decreased. The average size of egg-bearing Daphnia and Bosmina and the minimum size of egg-bearing specimens of the two genera rose. In the less eutrophic Lake Stigsholm, zooplankton and their grazing potential were also markedly affected by fish. However, the decrease in Chl a was slight, and phytoplankton biovolume, pH and the oxygen concentration were not affected. TN was higher when fish were absent. Our results indicate that: (i) there is a notable effect of fish on zooplankton community structure and size during winter in both eutrophic and hypertrophic North temperate lakes, (ii) Chl a can be high in winter in such lakes, despite low light irradiance, if fish are abundant, and (iii) the cascading effects on phytoplankton and nutrients in winter may be more pronounced in hypertrophic lakes. Climate warming supposedly leading to reduced winter mortality and dominance of small fish may enhance the risk of turbid state conditions in nutrient-enriched shallow lakes, not only during the summer season, but also during winter.     

Rapid Ecological Shift Following Piscivorous Fish Introduction to Increasingly Eutrophic and Warmer Lake Furnas (Azores Archipelago,Portugal): A Paleoecological Approach

Lake ecosystems are nowadays often subjected to multi-stressors, such as eutrophication, climate change, and fish manipulations, the effects of which can be difficult to disentangle, not least from the usual short-term limnological time-series that are available. However, multi-proxy paleoecological approaches may offer such opportunities, especially in the study of remote island lakes characterized by being species poor and buffered somewhat against the recent climate change. We used a paleoecological multi-proxy approach to determine the relative importance of nutrient loading, meteorological forcing, and fish species introduction for recent lake ecosystem development in Lake Furnas on the island of São Miguel, the Azores. The lake was stocked with cyprinids in the late nineteenth century and recently also with piscivorous fish, and has been affected by increasing agricultural activities in its catchment. We analyzed marker pigments, cladoceran remains, and subfossil diatoms in a 46-cm core representing the last 40 years. Remains of large-bodied cladocerans were virtually absent until the introductions of piscivorous pike and pikeperch in 1980 and 1982, respectively, after which the zooplankton community composition changed abruptly. First Ceriodaphnia sp. appeared (ca. 1980), followed by Daphnia a few years later. Carotenoids from cyanobacteria (myxoxanthophyll, aphanizophyll) were regularly present in the lake sediment with a major shift occurring around 1994, from N₂-fixing to non-fixing groups. This shift coincided with the onset of anoxia in the lake water, evidenced by the presence of pigments from Chlorobiaceae (BChl-e homologues and isorenieratene), and with a rapid decrease in benthic-tychoplanktonic diatoms and an increase in cyanobacteria, chlorophytes, dinoflagellates, and cryptophytes. The composition of microbial and algal assemblages changed rapidly after Daphnia appearance, and the covariance between fish stocking, nutrient loading, and enhanced temperatures captured most of the variability in algae accumulation, and thus likely in lake primary production as well. Thus, lake production apparently did not respond strongly to specific changes in temperature, food-web structure, or nutrient input, but rather to the combined effects of all the three forcing mechanisms, emphasizing the role of multi-stressors in lake ecosystem functioning. Our study demonstrates the sensitivity of these remote species-poor lakes to increased nutrient loading, introduction of non-native species, such as fish and climate change.     

Climate change effects on nitrogen loading from cultivated catchments in Europe: implications for nitrogen retention, ecological state of lakes and adaptation

Climate change might have profound effects on the nitrogen (N) dynamics in the cultivated landscape as well as on N transport in streams and the eutrophication of lakes. N loading from land to streams is expected to increase in North European temperate lakes due to higher winter rainfall and changes in cropping patterns. Scenario (IPCC, A2) analyses using a number of models of various complexity for Danish streams and lakes suggest an increase in runoff and N transport on an annual basis (higher during winter and typically lower during summer) in streams, a slight increase in N concentrations in streams despite higher losses in riparian wetlands, higher absolute retention of N in lakes (but not as percentage of loading), but only minor changes in lake water concentrations. However, when taking into account also a predicted higher temperature there is a risk of higher frequency and abundance of potentially toxic cyanobacteria in lakes and they may stay longer during the season. Somewhat higher risk of loss of submerged macrophytes at increased N and phosphorus (P) loading and a shift to dominance of small-sized fish preying upon the key grazers on phytoplankton may also enhance the risk of lake shifts from clear to turbid in a warmer North European temperate climate. However, it must be emphasised that the prediction of N transport and thus effects is uncertain as the prediction of regional precipitation and changes in land-use is uncertain. By contrast, N loading is expected to decline in warm temperate and arid climates. However, in warm arid lakes much higher N concentrations are currently observed despite reduced external loading. This is due to increased evapotranspiration leading to higher nutrient concentrations in the remaining water, but may also reflect a low-oxygen induced reduction of nitrification. Therefore, the critical N as well as P loading for good ecological state in lakes likely has to be lower in a future warmer climate in both north temperate and Mediterranean lakes. To obtain this objective, adaptation measures are required. In both climate zones the obvious methods are to change agricultural practices for reducing the loss of nutrients to surface waters, to improve sewage treatment and to reduce the storm-water nutrient runoff. In north temperate zones adaptations may also include re-establishment of artificial and natural wetlands, introduction of riparian buffer zones and re-meandering of channelised streams, which may all have a large impact on, not least, the N loading of lakes. In the arid zone, also restrictions on human use of water are urgently needed, not least on the quantity of water used for irrigation purposes.