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.     

Night,day, sunrise,sunset: do fish under snow and ice recognize the difference?

1. Although boreal lakes are ice‐covered for several months annually, little is known about the behaviour of fish under ice. To consider the reasons for diel vertical migrations (DVM) it is important to compare periods under ice as opposed to under open water. Echosounding provides a tool for non‐intrusive continuous monitoring of fish, even in winter. 2. Changes in the vertical distribution of fish through six 48‐h periods were monitored using a stationary, mounted echosounder that beamed vertically either from the bottom up or from the surface down from February to April, 2003. The up‐beaming and down‐beaming transducers were run alternately for 24 h each over the 48‐h period. Standard echo analysis software was used to detect fish traces and estimate the vertical and temporal distribution of fish abundance. Fish were sampled with a winter seine. 3. Prominent diel vertical migration in response to changing light level was detected throughout the study period (late winter to spring). Fish were highest in the water column at sunset and sunrise. In daylight, most detected fish were well below 10‐m depth. The number of fish detected was greatest during the night when they occurred throughout almost the whole water column, sometimes with a considerable number very close to the ice. Fish were mostly vendace and whitefish. 4. It became evident from data from the up‐beaming transducer that at night fish may occupy the layer closest to the surface. These fish would not have been detected if we had only used the down‐beaming transducer. The overall pattern of DVM in winter was very similar to that in summer. The results support the suggestion that DVM is a genetically fixed behavioural trait responding to the contemporary level of illumination.     

Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

Effects of habitat complexity on community structure and predator avoidance behaviour of littoral zooplankton in temperate versus subtropical shallow lakes

1. Structural complexity may stabilise predator–prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large‐bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N). We used plants of different architecture (submerged and free‐floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon‐richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger‐bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small‐bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant‐associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5‐fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plant‐associated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the free‐floating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predator‐avoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.     

Diverse Responses of Planktonic Crustaceans to Fish Predation by Shifts in Depth Selection and Size at Maturity

We investigated the behaviour and life histories of large zooplankton in the Esch-sur-Sûre reservoir (Luxembourg). We found that the decrease in size at maturity, as well as diurnal refuge in deep waters, were the adaptive responses (concurrently or alternatively) adopted by large zooplankters to cope with the increasing predation risk throughout the summer. Daphnia galeata initiated a more severe trade-off to cope with high summer predation relative to other cladoceran species. Diaphanosoma brachyurum and Daphnia cucullata seem less susceptible to fish predation as indicated by the low alteration of their size structure and their vertical distribution. The copepod Eudiaptomus gracilis took refuge in deep waters and achieved important modifications of its mature size.     

Diurnal changes in the vertical distribution of cladocerans in a biomanipulated lake

The diel vertical migration of cladoceran zooplankton in LakeVesijärvi was studied after a distinct improvement of the waterquality following mass removal of fish. Four out of seven speciesshowed diurnal changes in vertical distribution. The two speciesdominating at 0–10 m, Bosmina crassicornis and Daphniacucullata, migrated reverse, while hypolimnetic Bosminalongispina and Ceriodaphnia quadrangula ascended at night.The migration did not cross the thermocline, suggesting thatzooplankton had a minor role in restricting the availability ofphosphorus for phytoplankton. The reverse migration in theepilimnion was likely connected to the feeding behaviour of thedominating planktivores. During night, smelt (Osmeruseperlanus (L.)) migrated from the hypolimnion to the surface tofeed on daphnids, while roach (Rutilus rutilus (L.)) thatinhabited the 5–15 m depth, was day-active and fed mainly on Bosmina. In the hypolimnion, the cladocerans used the low oxygenconcentration as a refuge and at night migrated to layers moreprofitable for feeding. This suggests that the predation by fishwas the ultimate factor of the vertical migration, but otherenvironmental factors determined its magnitude.     

浮游动物昼夜垂直迁移机理的主要假说及其研究进展

对有关浮游动物昼夜垂直迁移(DVM)机理的实验、假说以及理论模型方面的研究进.展进行了综述。昼夜垂直迁移通常指常规迁移(傍晚上升,拂晓下降),其行为过程不仅影响浮游动物的垂直分布,而且也间接地影响其水平分布及生活史特征,对浮游动物在一定水域的种群维持和补充具有重要的意义。与垂直迁移机理有关的假说主要有光驱动假说、逃避捕食者假说、能量和资源利用假说等,其中逃避捕食者假说已得到大量的实验证实。其理论模型研究一般在两个时间尺度上进行,一个是短期的行为机制研究,另一个从长期的生活史策略的角度考虑。随着实验技术的进步和理论及建模工具的发展,这两个时间尺度的研究正逐渐统一到一个框架下进行。     

Variability and possible adaptive significance of day-time vertical distribution ofLeptodora kindtii (Focke) (Cladocera) in a shallow eutrophic lake

The day-time vertical distribution of different size classes ofLeptodora kindtii in Tjeukemeer was studied during two contrasting years: 1978 and 1979. During the first year (1978) the predation pressure exerted by the planktivorous fish was rather high, whilst this was ca. 15 times lower during 1979. The diel vertical migration of different size classes ofLeptodora, was studied twice during 1979. Both studies support the predator avoidance hypothesis and also emphasize the idea that diel vertical migration is a dynamic rather than a fixed behaviour trait within a population.     

Preliminary experiments on resource competition between a migrating and a non-migrating clone of the hybrid D. galeata×hyalina

Competition experiments between two clones of thehybrid D. galeata × hyalina differing invertical migration behaviour were carried out in thelaboratory, in a set-up which separated the clonesspatially. The influence of fish kairomones,artificial predation and the imitation of dielvertical migration circumstances were investigated. Inthe presence of fish kairomones, the non-migratingclone outnumbered the migrating one. The introductionof artificial predation reduced this effect in such away that the migrating clone had a competitiveadvantage over the non-migrating clone. It wasconcluded that competition results do not contradictlife history studies performed with the same twoclones, since the presence of fish kairomones enhancesthe reproduction of the non-migrating clone more thanthat of the migrating one.     

Quantification of diel vertical migration by micronektonic taxa in the northeast Atlantic

Analyses of day/night changes in the bathymetric distribution of micronektonic biomass at 16 stations in the northeastern Atlantic, sampled between 1978 and 1994, provided quantitative estimates of the organic carbon fluxes associated with diel vertical migration of individual taxa of micronekton. Gelatinous taxa contributed 50–80% of the integrated standing crop by volume but, apart from tunicates, contributed relatively little to the active migratory fluxes when expressed in terms of carbon. Total micronektonic migratory fluxes into the upper 200 m ranged from 12.5 to 58 mgC per m². At 15 stations, fish and pteropods provided 50–80% of the fluxes into the upper 100, 200 and 400 m. At one station, tunicates (pyrosomes) contributed substantially. Wherever tunicates or the medusa Pelagia were swarming, migrations by other taxa appear to be suppressed. The mean proportions of the stock (in terms of biomass) of each of the dominant migratory taxa entering and leaving the upper 100 m were 23% for tunicates, 18% for fish, 22% for pteropods, 8% for decapod crustaceans and 23% for euphausiids. The maximum proportions for these five taxa were 90%, 60%, 75%, 25% and 75%, respectively. Similar estimates of the mean fluxes into and out of the upper 400 m were generally higher: 19% for tunicates, 39% for fish, 28% for pteropods, 49% for decapods and 55% for euphausiids; the respective maxima were 99%, 74%, 99%, 72% and 91%. It is estimated for fish that if these migrations occur throughout the year, they will result in an active carbon export (both POC and DOC) from the wind-mixed layer and immediate sub-thermocline depths of about 500% of the mean annual standing stock. If this estimate can be extended to other taxa, then the material fluxes resulting from these active migrations will be quantitatively similar to those resulting from the deposition of phytodetritus at temperate latitudes.     

Population ecology, vertical migration and feeding of the Ponto-Caspian invader Hemimysis anomala in a gravel-pit lake connected to the River Rhine

1. The Ponto‐Caspian invader, Hemimysis anomala, was recently found in large numbers in a gravel‐pit lake connected to the Lower Rhine. Mysids were sampled with Perspex traps between September 2002 and April 2003 to study the population dynamics, vertical migration and feeding. 2. The abundance (as catch per unit effort, CPUE) of H. anomala declined from 270 individuals (ind.) trap^?1 (4 h)^?1 in December to below 4 ind. trap^?1 (4 h)^?1 in April. Average lengths ranged from 4.9 mm in autumn to 9.9 mm in March. The length–weight relationship of virgin females changed throughout the sampling period, from a size‐corrected wet weight of 5.1 mg in September to 16.6 mg in April for a female of 7.8 mm. 3. Successive monthly samples taken over 24 h revealed that H. anomala preferred the surface at twilight and night. During dawn the mysids migrated to the middle and bottom layers and were hardly found during day. Diel vertical migration clearly depends on the proximate factor light. 4. Stomach analyses revealed that larger H. anomala preferred zooplankton, whereas small individuals fed more on phytoplankton. The seasonal comparison showed an increasing percentage of zooplankton with increasing length. The proportion of zooplankton in the stomachs of large H. anomala individuals was highest during night and lowest during day. 5. The results suggest that H. anomala may become an important link between primary/secondary production and higher trophic levels in the food web of its new environments at the Lower Rhine.