Diel vertical migration in zooplankton: rapid individual response to predators

While diel vertical migration in zooplankton has been shownrecently to be a predator avoidance behavior, the mechanismby which predators induce and maintain such behavior has beendebated. We report results of an in situ predator manipulationexperiment during which enclosed populations of the marine planktomccopepod Acaraa hudsonica rapidly changed their vertical distributionand diel migration behavior depending on presence or absenceof the planktivorous fish Casterosteus aculeatus These resultspoint unambiguously to phenotypic behavioral plasticity of individualplanktonic prey, not, as previously hypothesized, population-geneticlevel behavioral changes caused by selective fish predation,as the mechanism underlying changes in diel vertical migrationin this copepod.     

Diel vertical migration arising in a habitat selection game

Predator and prey react to each other, adjusting their behavior to maximize their fitness and optimizing their food intake while keeping their predation risk as low as possible. In a pelagic environment, prey reduce their predation mortality by adopting a diel vertical migration (DVM) strategy, avoiding their predator during their peak performance by finding refuge in deep layers during daylight hours and feeding at the surface during the night. Due to the duality of the interaction between prey and predator, we used a game theory approach to investigate whether DVM can be a suitable strategy for the predator as well as the prey. We formulated three scenarios in plankton ecology in order to address this question. A novel finding is that mixed strategies emerge as optimal over a range of the parameter space, where part of the predator or prey population adopts a DVM while the rest adopt one or other “sit and wait” strategies.     

Diel vertical migration of zooplankton in the Northeast Atlantic

Acoustic Doppler current profiler (ADCP) data collected duringAugust–September 1991 reveal the diel migration of zooplanktonin the northeast Atlantic (50–60     

Diel vertical migration in Mesocyclops edax: implications for predation rate estimates

The diel migration patterns of Mesocyclops edax and its preyin a small lake were followed in two studies separated by approximatelyone year. Gut contents of the predators were examined and selectivityindices calculated at each depth at 0100 h during 1980. Thethree principal zooplankton prey found in the guts of M. edaxwere Keratella, Kellicottia, and Bosmina. The predator and allthree major prey species exhibited unique and different dielvertical distribution and migration patterns. The complex natureof the spatio-temporal variation in prey density to which M.edax is exposed, demonstrates the dangers of using selectivityindices without knowledge of the distribution patterns of bothpredator and prey. An increase in vertebrate predation pressurefrom one year to the next is thought to be responsible for anincrease in the abundance of small zooplankton species, thedisappearance of two out of three of the large zooplankton species,and the onset of a pronounced nocturnal migration pattern inthe third large species. ¹Present address: Biology Department, Williams Hall #31, LehighUniversity, Bethlehem, PA 18015, USA     

Diel vertical migration of Arctic zooplankton during the polar night

High-latitude environments show extreme seasonal variation in physical and biological variables. The classic paradigm of Arctic marine ecosystems holds that most biological processes slow down or cease during the polar night. One key process that is generally assumed to cease during winter is diel vertical migration (DVM) of zooplankton. DVM constitutes the largest synchronized movement of biomass on the planet, and is of paramount importance for marine ecosystem function and carbon cycling. Here we present acoustic data that demonstrate a synchronized DVM behaviour of zooplankton that continues throughout the Arctic winter, in both open and ice-covered waters. We argue that even during the polar night, DVM is regulated by diel variations in solar and lunar illumination, which are at intensities far below the threshold of human perception. We also demonstrate that winter DVM is stronger in open waters compared with ice-covered waters. This suggests that the biologically mediated vertical flux of carbon will increase if there is a continued retreat of the Arctic winter sea ice cover.     

Diel vertical migration ofEudiaptomus gracilis during a short summer period

Several aspects of a diel vertical migration (DVM) of adultEudiaptomus gracilis in Lake Maarsseveen (The Netherlands) are described. The period of DVM lasted from the end of May until the middle of August. On May 21, 1989, the population was found divided into a deep dwelling part and a part in the upper five meter. Large shoals of juvenile perch were observed in the open water for the first time. On June 7, the whole population was down below 10 m and concentrated in a zone of high chlorphyll-a concentrations. One week later, a regular DVM was performed. The amplitude of this migration gradually decreased towards the end of the migration period. The ascent in the evening and the descent in the morning took place after sunset and before sunrise, respectively. The movements coincided with high relative changes in light intensity. Population size increased rapidly during the period of DVM but decreased again before the end of this period.     

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.     

Diel vertical migration of adult Pacific cod Gadus macrocephalus in Alaska

The diel vertical migration (DVM) of Pacific cod Gadus macrocephalus was examined using depth and temperature data from 250 recaptured archival tags deployed on G. macrocephalus in the eastern Bering Sea and in the Gulf of Alaska near Kodiak Island. DVM of two types, deeper during daytime (type I) and deeper during night‐time (type II), occurred frequently (15–40% of all days) in G. macrocephalus released at all sites. Most individuals displayed both diel types, with each type of behaviour lasting up to 58 contiguous days, and day and night depth differences averaging c. 8 m. Despite high among‐individual variability, the occurrence of DVM varied significantly with the release site, season (i.e. day‐of‐year) and bottom depth, with the trend in seasonal occurrence nearly opposite for type I compared to type II DVM. No significance could be attributed to G. macrocephalus fork length, sex or ambient (tag) temperature. Trends in the magnitude of G. macrocephalus depth change were observed, with increased movement often occurring during night‐time, dawn and dusk, and at release sites where the bathymetry was more complex. Both type I and type II DVMs were attributed to foraging on prey species that also undergo DVM, and increased vertical movements of G. macrocephalus during crepuscular and night‐time periods were attributed to more active foraging during dim‐light conditions when G. macrocephalus can potentially exploit a sensory advantage over some of their prey.     

巢湖微囊藻和浮游甲壳动物昼夜垂直迁移的初步研究

2002年10月进行了巢湖微囊藻和几种优势浮游甲壳动物的昼夜垂直变化的研究,结果表明:微囊藻具有明显的昼夜垂直变化现象。白天上层水中的微囊藻密度显著高于下层水中,夜晚逐渐下沉使得下层水中的密度相对高于上层水。微囊藻与叶绿素a、水温、溶解氧和pH等均呈显著的正相关(p<0.01)。几种优势浮游甲壳动物的昼夜垂直迁移存在较大的差异。短尾秀体溞和角突网纹溞白天在下层水(1.5m和2.5m)中的密度较高,夜晚则倾向于在上层水(0m和0.5m)中活动。相反,卵形盘肠溞白天在上层水中密度较高,象鼻溞则在11:00和15:00时各水层中的密度显著高于夜晚。汤匙华哲水蚤和广布中剑水蚤白天倾向于在下层水中活动,夜晚则逐渐迁移到上层水中。许水蚤在夜晚和凌晨3:00时各水层中的密度显著高于白天。中华窄腹剑水蚤昼夜垂直变化不明显。微囊藻与短尾秀体溞密度呈显著的负相关,而与象鼻溞和卵形盘肠溞呈显著的正相关(p<0.01)。     

Diel vertical migration of zooplankton following optimal food intake under predation

A model is developed to investigate the trade-offs between benefitsand costs involved in zooplanktonic diel vertical migration(DVM) strategies. The ‘venturous revenue’ (VR) isused as the criterion for optimal trade-offs. It is a functionof environmental factors and the age of zooplankter. Duringvertical migration, animals are assumed to check instantaneouslythe variations of environmental parameters and thereby selectthe optimal behavioral strategy to maximize the value of VR,i.e. taking up as much food as possible with a certain riskof mortality. The model is run on a diel time scale (24 h) infour possible scenarios during the animal’s life history.The results show that zooplankton can perform normal DVM balancingoptimal food intake against predation risk, with the profileof DVM largely modified by the age of zooplankter.     

Control mechanisms of diel vertical migration: theoretical assumptions

We explore control mechanisms underlying the vertical migration of zooplankton in the water column under the predator-avoidance hypothesis. Two groups of assumptions in which the organisms are assumed to migrate vertically in order to minimize realized or effective predation pressure (type-I) and to minimize changes in realized or effective predation pressure (type-II), respectively, are investigated. Realized predation pressure is defined as the product of light intensity and relative predation abundance and the part of realized predation pressure that really affects organisms is termed as effective predation pressure. Although both types of assumptions can lead to the migration of zooplankton to avoid the mortality from predators, only the mechanisms based on type-II assumptions permit zooplankton to undergo a normal diel vertical migration (morning descent and evening ascent). The assumption of minimizing changes in realized predation pressure is based on consideration of DVM induction only by light intensity and predators. The assumption of minimizing changes in effective predation pressure takes into account, apart from light and predators also the effects of food and temperature. The latter assumption results in the same expression of migration velocity as the former one when both food and temperature are constant over water depth. A significant characteristic of the two type-II assumptions is that the relative change in light intensity plays a primary role in determining the migration velocity. The photoresponse is modified by other environmental variables: predation pressure, food and temperature. Both light and predation pressure are necessary for organisms to undertake DVM. We analyse the effect of each single variable. The modification of the phototaxis of migratory organisms depends on the vertical distribution of these variables.     

Diel vertical migration thresholds of Karenia brevis (Dinophyceae)

Light and nutrient availability change throughout dinoflagellate diel vertical migration (DVM) and/or with sub-population location in the water column along the west Florida shelf. Typically, the vertical depth of the shelf is greater than the distance a sub-population can vertically migrate during a diel cycle, limiting the ability of a sub-population to photosynthetically fix carbon toward the surface and access nutrients sub-surface. This project investigated changes of Karenia brevis (C.C. Davis) G. Hansen et Moestrup intracellular carbon, nitrogen, internal nitrate (iNO₃), free amino acid (FAA), and total lipid concentrations in high-light, nitrate-replete (960 μmol quanta m⁻² s⁻¹, 80 μM NO₃), and high-light, nitrate-reduced (960 μmol quanta m⁻² s⁻¹, <5 μM NO₃) mesocosms. The nitrate-reduced mesocosm had a slowed cell division rate when compared to the nitrate-replete mesocosm. Minimum intracellular carbon, nitrogen, iNO₃, FAA, and total lipid concentrations during the largest surface sub-population aggregations led to the conclusion that daughter cells resulting from cell division received unequal shares of the parental resources and that this inequality influenced migration behavior. Nutrient reduced daughter cells were more strongly influenced by light and phototaxis for carbon production than their replete same cell division sister cells during vertical migration thus rapidly increasing the fulfillment of constituents through photosynthesis. Vertical migration was consistent with an optimization scheme based on threshold limits through utilization or formation of photosynthate. We propose a simplified conceptual model describing how K. brevis is transported along the benthos of the west Florida shelf from off-shore to on-shore. Dynamic carbon thresholds are also suggested for future DVM modeling efforts on K. brevis populations transported between nitrogen replete and nitrogen reduced environmental conditions.     

Diel vertical migration of major fish-species in Lake Victoria, East Africa

Understanding of migration patterns is essential in the interpretation of hydro-acoustic stock assessment data of partly demersal partly pelagic fish stocks. In this paper we provide this kind of information for some species that were common in the Mwanza Gulf of Lake Victoria in the 1980s, before and after the upsurge of introduced Nile perch (Lates niloticus). Detritivorous haplochromines and Nile tilapia (Oreochromis niloticus), both stay near the bottom during day and night. Feeding seems to occur predominantly during the day. The zooplanktivorous haplochromines and dagaa (Rastrineobola argentea) dwell near the bottom by day and migrate towards the surface during the night. They seem to follow their prey, zooplankton and lake-fly larvae. Piscivorous nembe (Schilbe intermedius) show similar migration patterns to zooplanktivorous fishes, but their behaviour cannot be unambiguously explained by pursuit of prey. Nile perch to some extend migrate into the column at night, though the majority remains near the bottom. Feeding takes place during day and night.     

A modelling exploration of vertical migration by phytoplankton

The behaviour of phytoplankton having different abilities to assimilate N in darkness was considered in simulations of vertical migrations. Such behaviour is especially important for the competitive advantage of flagellates, including harmful algal species. Three phases of biomass development were apparent. (1) Cells remained at a subsurface location with migration down to avoid photoinhibitory light at midday; as the attenuation of light increased with biomass growth, the mean depth became shallower. (2) On exhaustion of nutrients in surface waters, cells migrated down through the nutricline in the latter half of the daylight period, with a subsurface maximum in the photic zone as long as light penetration matched requirements. When that condition was no longer met (3), cells migrated between the very surface (forming dense aggregations) and the nutricline. While the ability to perform dark N-assimilation is not critical when N-sources are available at low concentrations, it is important when (as encountered following migration down to a nutricline), nutrients are available at higher concentration in darkness. The most advantageous configuration tested, where nitrate assimilation (as well as that of ammonium) continued at a high rate in darkness as long as C-reserves remained, is not actually used in migratory species but in non-migratory diatoms. The use of the outwardly inferior configurations typical of migratory species, in which dark nitrate-assimilation is notably poorer than assimilation in the light, reflects a deficient metabolism or indicates that N-sources other than nitrate are more important. It is unfortunate then that most attention has been paid to nitrate nutrition in experiments on migrating species. While an ability to continue N-assimilation in darkness as well as during daylight is advantageous, there is no evidence for phytoplankton to be able to grow at high growth rates when decoupling photosynthesis at the surface and N-assimilation at depth.     

Diel vertical migration of a pelagic amphipod in the absence of fish predation

The pelagic amphipod, Hyalella montezuma, migrates vertically into the surface waters at twilight in Montezuma Well, Arizona, USA despite the absence of fish predators or thermal stratification. We suggest that a persistent, dense, neustonic algal assemblage may provide a food resource incentive for the twilight ascent.     

Diel vertical migration of copepods and its environmental drivers in subtropical Bahamian blue holes

Aquatic Ecology - Diel vertical migration (DVM) is the most common behavioral phenomenon in zooplankton, and numerous studies have evaluated DVM under strong seasonality at higher latitudes. Yet,...     

Diel vertical migration and feeding of chaetognaths in coastal waters of the eastern Mediterranean

This study investigates the diel vertical distribution and the diet of the most important chaetognath species found in the 0–50 m surface layer of a coastal area in the eastern Mediterranean during a 24-hour period in July 2004. Among the recorded chaetognaths, Sagitta enflata was the most abundant species (41.6%), followed by S. minima (32.5%) and S. serratodentata (20.8%). Those three species exhibited a “twilight migration” pattern, with only small differences among them. Vertical separation was found between S. enflata and S. minima. Both species preyed mainly on cladocerans, although copepods were the most abundant group in the zooplankton assemblage. The chaetognath species followed partially the diel vertical migration of their prey. S. enflata showed high feeding intensity at different times in both day and night, while S. minima fed more intensively at midday (12:00) and at dusk (20:00), and S. serratodentata in the morning (08:00). It seems that in order to coexist in an area of low productivity the chaetognath species follow the basic ecological rules of space, time and food-type separation, in order to reduce the inter- and intra-specific competition. The high preference of S. minima and especially of S. enflata for the cladocerans made them probably the most important predators of cladocerans during summer.