Diel vertical migration of freshwater fishes – proximate triggers,ultimate causes and research perspectives

1. Diel vertical migrations (DVM) are typical for many cold‐water fish species such as Pacific salmons (Oncorhynchus spp.) and coregonids (Coregonus spp.) inhabiting deep lakes. A comprehensive recent overview of DVM in freshwater fish has not been available, however. 2. The main proximate trigger of DVM in freshwater fish is the diel change in light intensity, with declining illumination at dusk triggering the ascent and the increase at dawn triggering the descent. Additional proximate cues are hydrostatic pressure and water temperature, which may guide fish into particular water layers at night. 3. Ultimate causes of DVM encompass bioenergetics efficiency, feeding opportunities and predator avoidance. None of these factors alone can explain the DVM in all cases. Multi‐factorial hypotheses, such as the ‘antipredation window’ combined with the thermal niche hypothesis, are more likely to explain DVM. It is suggested that planktivorous fish move within a layer sufficiently well illuminated to capture zooplankton, but too dark for predators to feed upon the migrating fish. In complete darkness, fish seek layers with a temperature that optimises bioenergetics efficiency. The strength of each factor may differ from lake to lake, and hence system‐specific individual analyses are needed. 4. Mechanistic details that are still poorly explored are the costs of buoyancy regulation and migration, the critical light thresholds for feeding of planktivorous and piscivorous fish, and predator assessment by (and size‐dependent predation risk of) the prey fish. 5. A comprehensive understanding of the adaptive value of DVM can be attained only if the behaviour of individual fish within migrating populations is explicitly taken into account. Size, condition and reproductive value differ between individuals, suggesting that migrating populations should split into migrants and non‐migrants for whom the balance between mortality risk and growth rate can differ. There is increasing evidence for this type of partial DVM within populations. 6. Whereas patterns of DVM are well documented, the evolution of DVM is still only poorly understood. Because experimental approaches at realistic natural scales remain difficult, a combination of comprehensive data sets with modelling is likely to resolve the relative importance of different proximate and ultimate causes behind DVM in fish.     

Diel variation in the vertical distribution of fish and plankton in Lake Kinneret: a 24-h study of ecological overlap

Diel vertical migration (DVM) behaviour is a predator avoidance mechanism observed within many zooplankton species in the presence of zooplanktivorous fish. A 24-h survey was carried out in June 1998 to investigate diel variation in the vertical distribution of fish, zooplankton and phytoplankton (chlorophyll) in Lake Kinneret, Israel. Fish revealed diel variation in vertical distribution but had no spatial overlap with zooplankton, and consequently no apparent influence on zooplankton dispersal. Zooplankton revealed some diel variation in distribution being affected by thermocline and oxycline position and movement of the internal the internal seiche wave. Cyclopoid species closely follow the movement of the seiche wave implying that, due to their greater motility, they are following conditions that are suitable to them. The Cladocera species and small rotifers only partly, which may be part of their phototaxic behaviour. Physical forces like convection, horizontal and vertical forcing probably have a role in contributing to a homogeneous distribution of the plankton by preventing stratification or interfering with the more motile zooplankton which may be attempting to migrate.     

Diel vertical migration of zooplankton: optimum migrating schedule based on energy accumulation

Zooplankton perform diel vertical migration (DVM) to avoid predators at the upper water layer, but often stay in the upper water layer throughout the day seeking food in spite of the presence of predators. This difference in migrating behavior has been explained by differences in environmental conditions or genetic differences. We examined theoretically how nutritious conditions of zooplankton individuals relate to determining different migrating behavior. A simple optimization model, maximizing the population growth rate, demonstrates that zooplankton individuals change their migrating behavior depending on the amount of accumulated energy. Such energy accumulation and its investment in reproduction are repeated every reproductive cycle. Therefore, unless the reproductive cycle is synchronized among individuals, different migrating behaviors will be observed within a population even if no genetic differences exist. Our model demonstrates that such coexistence of the two migrating behaviors is possible in natural Daphnia populations, and suggests that internal conditions of zooplankton individuals may be important as a factor for determining migrating behavior of zooplankton. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cyclic temperatures influence growth efficiency and biochemical body composition of vertically migrating fish

1. Diel vertical migrations (DVM) are frequently observed in pelagic fish and zooplankton populations. In addition to predator avoidance and foraging opportunity, bioenergetics optimisation has been hypothesised to cause the selective advantage of migrating fish. However, experimental confirmation of growth advantages of fish held at naturally observed cyclic temperatures, and food densities are surprisingly rare. 2. We compared growth rates, growth efficiencies and energy budgets of vendace (Coregonus albula, Salmoniformes) fed daily rations of 10% body mass and held at low (4.5 °C), high (8 °C) and cyclic (switch between 4.5 and 8 °C) temperatures over a period of 6 weeks. Biochemical body composition was evaluated by bioimpedance analyses and direct determination of water, lipid and protein content in a subset of fish. 3. Growth rate and food conversion efficiency were similar in fish held at high and cyclic temperatures, but higher than those found at low temperatures. Body condition of fish at cyclic temperatures was maximised, but these fish also had the highest water content. The proportion of lipids and proteins was slightly depleted in fish held at cyclic and low temperatures relative to the high temperature treatment. Accordingly, growth and biochemical composition of fish responded specifically to cyclic temperatures and were not simply intermediate between those found at low and high temperatures. 4. We could not confirm a bioenergetics advantage of fish switching between high and low temperatures. However, there were no substantial extra metabolic costs through exposure to cyclic temperatures, and hence, fish performing DVM may benefit from predation avoidance without compromising their metabolic balance and hence growth rates. These results suggest that the evolution of DVM is a multi‐faceted process with no single ultimate explanation.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.     

From inducible defences to population dynamics: modelling refuge use and life history changes in Daphnia

We investigated the relative importance of a behavioural defence (refuge use through diel vertical migration) and a life history change (a reduced size at first reproduction) that are used by daphnids to decrease the risk of predation by visually hunting fish. We used an individual based model of a Daphnia population in a stratified lake to quantify the effects of these inducible defences on Daphnia predation-mortality and the resulting Daphnia population dynamics. Our analysis shows that diel vertical migration (DVM) confers a much stronger protection against fish predation than a reduced size at first reproduction (SFR). DVM allows daphnids to withstand a higher predation pressure in the epilimnion and it decelerates a Daphnia population decline more strongly than a reduced SFR. DVM effectively reduces the (P/B) flow of carbon from daphnids to fish.
Many theoretical studies have only considered the fitness benefits of DVM above 'staying up' in the epilimnion of a lake. Our results suggest that 'staying down' in the hypolimnion would confer an even stronger fitness benefit to Daphnia than DVM at times of peak predation risk. Daphnids that remain in the hypolimnion avoid the predation suffered by migrating daphnids around dusk and dawn. Staying down could prevent a Daphnia population decline, while DVM and a reduced SFR can only decelerate the decrease of Daphnia population densities under heavy fish predation. Staying down at high concentrations of fish infochemicals has in fact been observed within a variety of Daphnia clones and species, both in the laboratory and in stratified lakes.     

Blooms of Mesodinium rubrum in Southampton Water--do they shape mesozooplankton distribution?

Blooms of the red-tide ciliate Mesodinium rubrum occur everyyear in Southampton Water from late May to August, peaking inabundance in July. During blooms, M.rubrum concentrates in subsurfacelayers during the day and and also undergoes diurnal verticalmigration; this results in supersaturation in the near-surfacewater and significant deoxygenation at lower depths. Duringthe period of M.rubrum bloom, the mesozooplankton communityof Southampton Water is typically dominated by nauplius larvaeand calanoid copepods. The true nature of M.rubrum's impacton mesozooplankton and fish communities is equivocal, with contrastingreports of either bloom-associated avoidance or mortality byzooplankton or fish, or both, or alternatively grazing by micro-and mesoplankton on M.rubrum. A temporal sequence of zooplanktonhauls conducted between mid-May and early August indicated thatvertical profiles of mesozooplankton composition and abundancetypically exhibited lower near-surface densities, with numbersincreasing significantly below 3 m. During blooms, this patternoccurred both inside and outside patches, and therefore thereduced near-surface zooplankton numbers are considered to becaused primarily by turbulent mixing rather than by the associatedoxygen supersaturation induced by M.rubrum. Equally, no clearpattern of mesozooplankton density differences was observedat lower depths, associated with the low concentrations of oxygencaused by M.rubrum respiration. The results of the samplingprogramme undertaken show that M.rubrum does not significantlyexclude zooplankton from the vicinity of patches, nor is thereany associated substantial increase in zooplankton numbers.It is concluded that M.rubrum blooms have no immediate significantimpact on the broad structure of the mesozooplankton communitywithin Southampton Water.     

Seasonal vertical distribution and diel migration of zooplankton in a temperate stratified lake

The investigation of the vertical distribution of the zooplankton community in the temperate Lake Trichonis during four seasons in 2005, showed the existence of vertical segregation among species, ontogenetic stages and sexes within and between the major groups. In each season, the two or three more abundant rotifer species distributed at separate depth layers, while this feature was maintained during the entire 24 h period, since no diel vertical migrations (DVM) were performed. In contrast, the crustacean community, comprised mainly by the calanoid copepod Eudiaptomus drieschi and the cladoceran Diaphanosoma orghidani, showed various patterns of DVM, being more pronounced in spring and summer. Females of E. drieschi distributed deeper than males, while the copepod nauplii were found mainly in the surface layer in all four seasons. Temperature was the most important abiotic factor affecting directly and indirectly the vertical distribution and migration of various species. During stratification, the metalimnion was the most productive layer in Lake Trichonis, having maximum values of dissolved oxygen and low transparency due to high concentration of organic matter and phytoplankton. The DVM patterns of the crustaceans indicate that the metalimnion acts probably as a daylight refuge against predation by Atherina boyeri, which is the dominant planktivorous fish in the lake.     

On the cost of vertical migration: are feeding conditions really worse at greater depths?

SUMMARY 1. The ultimate explanation for diel vertical migration (DVM) of zooplankton is the avoidance of visual predation in surface waters. Studies on migrating zooplankton have shown that remaining in the cold and food-poor hypolimnion during the day, however, has demographic costs. Higher temperatures and greater food concentrations in the surface waters are thought to be the main reasons why Daphnia species move upwards at night.
2. In this study, we investigated the growth condition of daphniids raised on seston taken from different depths from a lake with and without a deep-water chlorophyll maximum.
3. Juvenile growth rates of Daphnia galeata x hyalina from the lake without a deep-water chlorophyll maximum were similar for all treatments. After temperature correction, however, growth rates were significantly higher on seston taken from the surface layers.
4. In contrast, in the lake with the deep-water chlorophyll maximum, D. galeata growth rates were higher in deeper strata, even after temperature correction. Although this lake had a weak temperature gradient, D. galeata left the food-rich strata at night and migrated into the surface food-poor environment. Invertebrate predation and oxygen depletion are probably not the reasons for the nocturnal upward migration into the surface strata. Therefore, we assume that D. galeata migrates upwards to take advantage of higher temperatures. Using several temperature–egg-development models, we could not, however, fully explain this behaviour.     

Alteration of photoresponses involved in diel vertical migration of a crab larva by fish mucus and degradation products of mucopolysaccharides

Photoresponses involved in the descent phase of nocturnal diel vertical migration (DVM) of larvae of the crab Rhithropanopeus harrisii were measured in a laboratory system that mimicked the underwater angular light distribution. The test hypothesis was that kairomones from fish that activate zooplankton photoresponses involved in DVM are derived from polysaccharides from the external mucus of fishes. Studies considered fish mucus from the mummichog (Fundulus heteroclitus) and disaccharides (originating from chondroitin sulfate A and heparin polysaccharides) that are likely constituents of fish mucus. R. harrisii larvae descend at sunrise with an isolume and remain near the isolume during the day. Since depth maintenance near the isolume depends upon a negative phototaxis, the lowest light intensity (threshold) that induces this response was used to quantify the effects of the test chemicals. It was predicted that exposure to fish kairomones would lower the photoresponse threshold, thereby resulting in larvae remaining deeper in the water column where light for visual predation was reduced. The photoresponse threshold declined as the concentration of fish mucus increased. Disaccharides originating from chondroitin sulfate A and heparin also decreased the photoresponse threshold as compared to responses in aged, filtered seawater. Collectively, the results support the hypothesis and indicate that disaccharide degradation products of predator mucus containing sulfated and acetylated amines can serve as kairomones.     

Hunt warm, rest cool: bioenergetic strategy underlying diel vertical migration of a benthic shark

1. Diel vertical migration (DVM) is a widespread phenomenon among marine and freshwater organisms and many studies with various taxa have sought to understand its adaptive significance. Among crustacean zooplankton and juveniles of some fish species DVM is accepted widely as an antipredator behaviour, but little is known about its adaptive value for relatively large-bodied, adult predatory fish such as sharks. Moreover, the majority of studies have focused on pelagic forms, which raises the question of whether DVM occurs in bottom-living predators. 2. To investigate DVM in benthic predatory fish in the marine environment and to determine why it might occur we tracked movements of adult male dogfish (Scyliorhinus canicula) by short- and long-term acoustic and archival telemetry. Movement studies were complemented with measurements of prey abundance and availability and thermal habitat within home ranges. A thermal choice experiment and energy budget modelling was used to investigate trade-offs between foraging and thermal habitat selection. 3. Male dogfish undertook normal DVM (nocturnal ascent) within relatively small home ranges (-100 x 100 m) comprising along-bottom movements up submarine slopes from deeper, colder waters occupied during the day into warmer, shallow prey-rich areas above the thermocline at night. Few daytime vertical movements occurred. Levels of activity were higher during the night above the thermocline compared to below it during the day indicating they foraged in warm water and rested in colder depths. 4. A thermal choice experiment using environmentally realistic temperatures supported the field observation that dogfish positively avoided warmer water even when it was associated with greater food availability. Males in laboratory aquaria moved into warm water from a cooler refuge only to obtain food, and after food consumption they preferred to rest and digest in cooler water. 5. Modelling of energy budgets under different realistic thermal-choice scenarios indicated dogfish adopting a 'hunt warm - rest cool' strategy could lower daily energy costs by just over 4%. Our results provide the first clear evidence that are consistent with the hypothesis that a benthic marine-fish predator utilizes DVM as an energy conservation strategy that increases bioenergetic efficiency.     

Temperature selection in Brook Charr: lab experiments,field studies,and matching the Fry curve

Zooplankton diel vertical migration (DVM) is an ecologically important process, affecting nutrient transport and trophic interactions. Available measurements of zooplankton displacement velocity during the DVM in the field are rare; therefore, it is not known which factors are key in driving this velocity. We measured the velocity of the migrating layer at sunset (upward bulk velocity) and sunrise (downwards velocity) in summer 2015 and 2016 in a lake using the backscatter strength (VBS) from an acoustic Doppler current profiler. We collected time series of temperature, relative change in light intensity chlorophyll-a concentration and zooplankton concentration. Our data show that upward velocities increased during the summer and were not enhanced by food, light intensity or by VBS, which is a proxy for zooplankton concentration and size. Upward velocities were strongly correlated with the water temperature in the migrating layer, suggesting that temperature could be a key factor controlling swimming activity. Downward velocities were constant, likely because Daphnia passively sink at sunrise, as suggested by our model of Daphnia sinking rate. Zooplankton migrations mediate trophic interactions and web food structure in pelagic ecosystems. An understanding of the potential environmental determinants of this behaviour is therefore essential to our knowledge of ecosystem functioning.

     

Two hundred years of zooplankton vertical migration research

Vertical migration is a geographically and taxonomically widespread behaviour among zooplankton that spans across diel and seasonal timescales. The shorter-term diel vertical migration (DVM) has a periodicity of up to 1 day and was first described by the French naturalist Georges Cuvier in 1817. In 1888, the German marine biologist Carl Chun described the longer-term seasonal vertical migration (SVM), which has a periodicity of ca. 1 year. The proximate control and adaptive significance of DVM have been extensively studied and are well understood. DVM is generally a behaviour controlled by ambient irradiance, which allows herbivorous zooplankton to feed in food-rich shallower waters during the night when light-dependent (visual) predation risk is minimal and take refuge in deeper, darker waters during daytime. However, DVMs of herbivorous zooplankton are followed by their predators, producing complex predator–prey patterns that may be traced across multiple trophic levels. In contrast to DVM, SVM research is relatively young and its causes and consequences are less well understood. During periods of seasonal environmental deterioration, SVM allows zooplankton to evacuate shallower waters seasonally and take refuge in deeper waters often in a state of dormancy. Both DVM and SVM play a significant role in the vertical transport of organic carbon to deeper waters (biological carbon sequestration), and hence in the buffering of global climate change. Although many animal migrations are expected to change under future climate scenarios, little is known about the potential implications of global climate change on zooplankton vertical migrations and its impact on the biological carbon sequestration process. Further, the combined influence of DVM and SVM in determining zooplankton fitness and maintenance of their horizontal (geographic) distributions is not well understood. The contrasting spatial (deep versus shallow) and temporal (diel versus seasonal) scales over which these two migrations occur lead to challenges in studying them at higher spatial, temporal and biological resolution and coverage. Extending the largely population-based vertical migration knowledge base to individual-based studies will be an important way forward. While tracking individual zooplankton in their natural habitats remains a major challenge, conducting trophic-scale, high-resolution, year-round studies that utilise emerging field sampling and observation techniques, molecular genetic tools and computational hardware and software will be the best solution to improve our understanding of zooplankton vertical migrations.     

Zooplankton biomass distribution patterns along the western Antarctic Peninsula (December 2002)

The phytoplankton [chlorophyll a (Chl a)], microzooplankton,mesozooplankton and macrozooplankton biomass and distributionwere studied as part of a multidisciplinary project (Tempano)along the Antarctic Peninsula during December 2002. Even thoughthe summer phytoplankton bloom was not yet developed in thearea, autotrophs dominated the plankton biomass. Phytoplanktonvertical distribution was, in general, homogeneous in the upper40–50 m of the water column, further decreasing with depth.Protozoans showed low biomass; their contribution to the totalplankton being one order of magnitude lower than that of autotrophs.The vertical distribution of protozoans was variable among stationswith marked peaks at depths ranging from 30 to 80 m. Mesozooplankton-integratedbiomass was generally low, although there was a notable increasesouthward near the ice marginal zone. Macrozooplankton distributionwas more variable without any clear zonal distribution pattern.The vertical distribution of meso- and macrozooplankton (>4mm) biomass showed clear peaks of abundance comprising differentspecies depending on the geographical area. Our biomass distributiondata suggest a food-web scenario in which macrozooplankton arepreying on mesozooplankton populations only in the northernerstations, and mesozooplankton are, in their turn, shaping theabundance of the emerging populations of microzooplankton. Phytoplankton,on the other hand, seem to be hardly controlled by grazing activity.     

Temperature sensitivity of vertical distributions of zooplankton and planktivorous fish in a stratified lake

Recent studies have indicated that temporal mismatches between interacting populations may be caused by consequences of global warming, for example rising spring temperatures. However, little is known about the impact of spatial temperature gradients, their vulnerability to global warming, and their importance for interacting populations. Here, we studied the vertical distribution of two planktivorous fish species (Coregonus spp.) and their zooplankton prey in the deep, oligotrophic Lake Stechlin (Germany). The night-time vertical centre of gravity both of the fish populations and of two of their prey groups, daphnids and copepods, were significantly correlated to the seasonally varying water temperature between March and December 2005. During the warmer months, fish and zooplankton occurred closer to the surface of the lake and experienced higher temperatures. The Coregonus populations differed significantly in their centre of gravity; hence, also, the temperature experienced by the populations was different. Likewise, daphnids and copepods occurred in different water depths and hence experienced different temperatures at least during the summer months. We conclude that any changes in the vertical temperature gradient of the lake as a result of potential future global warming may impact the two fish populations differently, and may shape interaction strength and timing between fish and their zooplankton prey. Priority programme of the German Research Foundation—contribution 9.     

The impact of diel vertical migration of <Emphasis Type="Italic">Daphnia</Emphasis> on phytoplankton dynamics

Diel vertical migration (DVM) of large zooplankton is a very common phenomenon in the pelagic zone of lakes and oceans. Although the underlying mechanisms of DVM are well understood, we lack experimental studies on the consequences of this behaviour for the zooplankton’s food resource—the phytoplankton. As large zooplankton species or individuals migrate downwards into lower and darker water strata by day and upwards into surface layers by night, a huge amount of herbivorous biomass moves through the water column twice a day. This migration must have profound consequences for the phytoplankton. It is generally assumed that migration supports an enhanced phytoplankton biomass and a change in the composition of the phytoplankton community towards smaller, edible algae in the epilimnion of a lake. We tested this assumption for the first time in field experiments by comparing phytoplankton biomass and community assemblage in mesocosms with and without artificially migrating natural stocks of Daphnia hyalina. We show that DVM can enhance phytoplankton biomass in the epilimnion and that it has a strong impact on the composition of a phytoplankton community leading to an advantage for small, edible algae. Our results support the idea that DVM of Daphnia can have strong effects on phytoplankton dynamics in a lake.     

Food resource effects on diel movements and body size of cisco in north-temperate lakes

The movement patterns and body size of fishes are influenced by a host of physical and biological conditions, including temperature and oxygen, prey densities and foraging potential, growth optimization, and predation risk. Our objectives were to (1) investigate variability in vertical movement patterns of cisco (Coregonus artedi) in a variety of inland lakes using hydroacoustics, (2) explore the causal mechanisms influencing movements through the use of temperature/oxygen, foraging, growth, and predation risk models, and (3) examine factors that may contribute to variations in cisco body size by considering all available information. Our results show that cisco vertical movements vary substantially, with different populations performing normal diel vertical migrations (DVM), no DVM, and reverse DVM in lakes throughout Minnesota and northern Wisconsin, USA. Cisco populations with the smallest body size were found in lakes with lower zooplankton densities. These smaller fish showed movements to areas of highest foraging or growth potential during the day and night, despite moving out of preferred temperature and oxygen conditions and into areas of highest predation risk. In lakes with higher zooplankton densities, cisco grew larger and had movements more consistent with behavioral thermoregulation and predator avoidance, while remaining in areas with less than maximum foraging and growth potential. Furthermore, the composition of potential prey items present in each lake was also important. Cisco that performed reverse DVM consumed mostly copepods and cladocerans, while cisco that exhibited normal DVM or no migration consumed proportionally more macro-zooplankton species. Overall, our results show previously undocumented variation in migration patterns of a fish species, the mechanisms underlying those movements, and the potential impact on their growth potential.     

Effects of Ultraviolet Radiation on Diel Vertical Migration of Crustacean Zooplankton: An in situ Mesocosm Experiment

The objective of this study was to expand the spatial scale of previous experiments on the effects of ultraviolet radiation (UVR) on diel vertical migration (DVM) by freshwater zooplankton. We conducted an in situ mesocosm experiment in highly UVR transparent Lake Giles, Pennsylvania, in which we imposed two treatments: ambient UVR and UVR-shielded. Mesocosms (3440 L, 0.74 m diameter, 8 m deep) were large enough to include a spatial refuge from UVR and permit relatively large-scale DVM. Daphnia catawba adopted a significantly deeper distribution during the day in the ambient UVR treatment compared to the UVR-shielded treatment, but effects of UVR were absent at night. In contrast, DVM by Leptodiaptomus minutus was unaffected by the UVR treatment. In both treatments, Leptodiaptomus minutus were most abundant at the bottom of the mesocosms during the day and exhibited a more uniform distribution across depths at night. These results suggest that UVR, along with temperature, algal resources, and predators, may affect zooplankton DVM in aquatic ecosystems.