Small-scale diel vertical migration of zooplankton in the High Arctic

Diel vertical migration (DVM) of zooplankton is considered less prominent at high latitudes where diel changes in irradiance are minimal during periods of midnight sun and polar night, leaving zooplankton without a temporal refuge and thus eliminating a key advantage of DVM. One of the shortcomings of previous DVM studies of zooplankton based on net sampling is that the depth resolution often has been too coarse to detect vertical migrations over short distances. We investigated DVM of zooplankton during August 2010 in drifting sea ice northeast of Svalbard (~81.5°N, ~30.5°E). Classical DVM behaviour (midnight rising, midday sinking) was observed between 20 and 80 m in young copepodite stages (CI–III) of Calanus finmarchicus and Calanus glacialis. The copepods Microcalanus spp., Pseudocalanus spp., Oithona atlantica, Oithona similis and Triconia borealis, alongside Eukrohnia hamata, Limacina helicina, and Fritillaria spp., all displayed signs of DVM. We conclude that zooplankton exhibit DVM in ice-covered waters over rather short distances to optimise food intake in the presence of predators.     

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.     

The effect of different zooplankton grazing patterns resulting from diel vertical migration on phytoplankton growth and composition: a laboratory experiment

Diel vertical migration (DVM) of herbivorous zooplankton is a widespread behavioural phenomenon in freshwater ecosystems. So far only little attention has been paid to the impact of DVM on the phytoplankton community in the epilimnion. Some theoretical models predict that algal population growth in the epilimnion should depend on the herbivores migration and grazing patterns: even if migrating zooplankton consume the same total amount of algae per day in the epilimnion as non-migrating zooplankton, nocturnal grazing should result in enhanced algal growth and favour algal species with high intrinsic growth rates over species with lower intrinsic growth rates. To test these hypotheses we performed experiments in which several algal species were confronted with different feeding regimes of Daphnia. In the experiments algal growth did not only depend on the absolute time of grazing but was comparatively higher when grazing took place only during the night, even when the grazing pressure was the same. Furthermore, algal species with higher intrinsic growth rates had higher advantages when being grazed upon only discontinuously during the night than algal species with a smaller intrinsic growth rate. The grazing pattern itself was an important factor for relative algal performance.     

Effects of moonlight on the vertical migration patterns of demersal zooplankton

The diel vertical migration patterns of demersal zooplankton, those organisms which habit bottom substrates but periodically emerge to swim freely in the water column, water determined throughout the lunar cycle. Demersal zooplankton were quantitatively sampled on a subtidal sand flat in the Gulf of California every 2 h for 24-h periods at new, full, first, and last-quarter moons, both as they emerged into the water column and as they returned to the benthos. Demersal zooplankton rarely migrated during daylight. Three general patterns of migration were observed. (1) Polychaetes and cumaceans emerged from the benthos at dusk, regardless of the phase of the moon. Polychaetes returned to the benthos throughout the night while cumaceans returned near dawn. (2) Species of amphipods and isopods exhibited significant avoidance of moonlight, delaying emergence until moonset or returning to the benthos at moonrise. (3) Species of copepods, mysids, shrimp, Branchiostoma (cephalochordate), and tanaids emerged into the water column throughout the night. The timing of migration was highly variable and did not correlate with the presence or absence of moonlight. Large zooplankton migrated less frequently into the water column during moonlit periods than small forms, suggesting that nocturnal predation by visually oriented planktivorous fish may be an important selective pressure.Demersal zooplankton emerged into artificially darkened emergence traps in significantly higher numbers during daylight and during full and quarter moons than into undarkened control traps, demonstrating that absence of light is a major cue stimulating migration. Reentry traps resting on the bottom captured higher densities of demersal zooplankton than either emergence traps or reentry traps suspended off the bottom. Thus, many demersal zooplankton remain near the bottom, rarely swimming far into the water column. Some trap avoidance was observed and current methods for collecting demersal zooplankton are evaluated. Since most demersal zooplankton remained in the water column only a short time, dispersal, particularly over short distances, may be a major advantage of migratory behavior. Migration facilitates rapid recolonization of disturbed or defaunated sites, disrupts and mixes bottom sediments, and results in daily variation in the microdistribution, patchiness, and species composition of the benthic fauna.     

The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton

Many pelagic animal species in the marine environment and in lakes migrate to deeper water layers before sunrise and return around sunset. The amplitude of these diel vertical migrations (DVM) varies from several hundreds of metres in the oceans to approx. 5–20 m in lakes. DVM can be studied from a proximate and an ultimate point of view. A proximate analysis is intended to reveal the underlying behavioural mechanism and the factors that cause the daily displacements. The ultimate analysis deals with the adaptive significance of DVM and the driving forces that were responsible for the selection of the traits essential to the behavioural mechanism. The freshwater cladoceran Daphnia is the best studied species and results can be used to model migration behaviour in general. Phototaxis in Daphnia spp., which is defined as a light-oriented swimming towards (positive phototaxis) or away (negative phototaxis) from a light source, is considered the most important mechanism basic to DVM. A distinction has been made between primary phototaxis which occurs when light intensity is constant, and secondary phototaxis which is caused by changes in light intensity. Both types of reaction are superimposed on normal swimming. This swimming of Daphnia spp. consists of alternating upwards and downwards displacements over small distances. An internal oscillator seems to be at the base of these alternations. Primary phototaxis is the result of a dominance of either the upwards or the downwards oscillator phase, and the direction depends on internal and external factors: for example, fish-mediated chemicals or kairomones induce a downwards drift. Adverse environmental factors may produce a persistent primary phototaxis. Rare clones of D. magna have been found that show also persistent positive or negative primary phototaxis and interbreeding of the two types produces intermediate progeny: thus a genetic component seems to be involved. Also secondary phototaxis is superimposed on normal swimming: a continuous increase in light intensity amplifies the downwards oscillator phase and decreases the upwards phase. A threshold must be succeeded which depends on the rate and the duration of the relative change in light intensity. The relation between both is given by the stimulus strength versus stimulus duration curve. An absolute threshold or rheobase exists, defined as the minimum rate of change causing a response if continued for an infinitely long time. DVM in a lake takes place during a period of 1-5-2 h when light changes are higher than the rheobase threshold. Accelerations in the rate of relative increase in light intensity strongly enhance downwards swimming in Daphnia spp. and this enhancement increases with increasing fish kairomone and food concentration. This phenomenon may represent a ‘decision-making mechanism’ to realize the adaptive goal of DVM: at high fish predator densities, thus high kairomone concentrations, and sufficiently high food concentrations, DVM is profitable but not so at low concentrations. Body axis orientation in Daphnia spp. is controlled with regard to light-dark boundaries or contrasts. Under water, contrasts are present at the boundaries of the illuminated circular window which results from the maximum angle of refraction at 48–9° with the normal (Snell's window). Contrasts are fixed by the compound eye and appropriate turning of the body axis orients the daphnid in an upwards or an obliquely downwards direction. A predisposition for a positively or negatively phototactic orientation seems to be the result of a disturbed balance of the two oscillators governing normal swimming. Some investigators have tried to study DVM at a laboratory scale during a 24 h cycle. To imitate nature, properties of a natural water column, such as a large temperature gradient, were compressed into a few cm. With appropriate light intensity changes, vertical distributions looking like DVM were obtained. The results can be explained by phototactic reactions and the artificial nature of the compressed environmental factors but do not compare with DVM in the field. A mechanistic model of DVM based on phototaxis is presented. Both, primary and secondary phototaxis is considered an extension of normal swimming. Using the light intensity changes of dawn and the differential enhancement of kairomones and food concentrations, amplitudes of DVM could be simulated comparable to those in a lake. The most important adaptive significance of DVM is avoidance of visual predators such as juvenile fish. However, in the absence of fish kairomones, small-scale DVMs are often present, which were probably evolved for UV-protection, and are realized by not enhanced phototaxis. In addition, the ‘decision-making mechanism’ was probably evolved as based on the enhanced phototactic reaction to accelerations in the rate of relative changes in light intensity and the presence of fish kairomones.     

Sampling and analysis of gut contents in relation to environmental variability and diel vertical migration by herbivorous zooplankton

The accuracy of standard sampling and analysis procedures forestimating ingestion by herbivorous zooplankton was assessedusing models. Artificial environments were created in a computermodel, allowing for depth-dependent variability in temperature,chlorophyll and primary production. Model zooplankton were simulatedwithin these artificial environments using individual-basedmodels. The model zooplankton feed and defaecate at rates determinedby temperature and food concentrations, and also exhibit dielvertical migration (DVM) according to a variety of migrationmodels. The computer model was run for different combinationsof these nine environmental and five DVM models. Data were ‘sampled’from the model output, similar to field sampling of mesozooplanktongrazing. Daily ingestion was calculated from the gut ‘samples’using standard procedures for analysing gut fluorescence. Thesample results were compared with the actual ingestion valuesin the model, and some causes of discrepancies were noted. (i)If incorrect temperatures were assumed when calculating thegut evacuation rate (K), then estimates of ingestion were wrongby up to 40%. (ii) Non-uniform food environments gave errorsof up to 30% because of the large variability of measured gutcontents among individuals. (iii) Sampling from only part ofthe total depth range (e.g. at the chlorophyll maximum) resultedin estimates of ingestion being only 5% of the real value. Thissampling practice should be discouraged, because the sampleis not random. (iv) If sampling is not frequent enough, errorscan be as large as 45%, but more usually were 10% for realisticsampling frequencies. We describe an analysis procedure thatuses Monte Carlo-type simulations in a computer spreadsheetto estimate population consumption. These calculations takeinto account natural variability due to populations, samplesand assumptions. We urge that results should be presented asranges of possible values, rather than as single ‘mean’values, to allow for easier recognition of meaningful differencesamong samples and systems.     

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.     

Small-scale distribution and diel vertical migration of zooplankton in a shallow lake (Lake Naardermeer,the Netherlands)

Groundwater is a major influence on the hydrological, chemical and thermal regime of chalk streams in the southern U.K. However, little is currently known about the nature of the sediment delivery system within these chalk stream systems, even though sediment-related problems have been increasingly cited as a cause of habitat degradation and of declining salmonid stocks. To address this knowledge gap, suspended sediment fluxes were monitored at 4 sites within the Hampshire Avon catchment between February 1999 and August 2000. Maximum suspended sediment concentrations ranged from nearly 45 mg l^–1 to 260 mg l^–1. Over the study period, annual suspended sediment loads ranged from 644 to 6215 t yr^–1 and annual specific sediment yields ranged from 1.4 to 12.5 t km^–2 yr^–1. The results show that, relative to other U.K. rivers, the study chalk streams are characterised by low suspended sediment concentrations and loads and less episodic behaviour.     

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     

Laboratory models of diel vertical migration in the dinoflagellate Ceratium hirundinella

SUMMARY. Diel vertical migrations of a dinoflagellate, Ceratium hirundinella , were induced in a laboratory tube (1.63 × 0.15 m) under a light-dark cycle. The timing of vertical migrations differed between cultures in the exponential and stationary phases of growth; the latter showed a greater coincidence with the light regime.
Migration of cells into the surface layers occurred at low values of surface irradiance (<550 μeinsteins m⁻² s⁻¹). At irradiances more closely approaching summer sunshine (> 1300 μE m⁻² s⁻¹) there was a marked avoidance of surface waters, and population maxima were found at depths associated with a relative irradiance level of 10% or c. 150 μE m⁻² s⁻¹). Thermal stratification restricted downward movement of cells into the cooler layers. The combination of high surface irradiance and thermal stratification resulted in large, stable, sub-surface maxima of Ceratium , similar to those observed in natural waters under comparable environmental conditions.     

Fluorescence analysis of zooplankton gut contents and an investigation of diel feeding patterns

A method is described for measurement of the in situ feeding rate of planktonic herbivores. Freshly caught Zooplankton are washed with filtered sea water and frozen. The gut contents are later analysed using acetone extracts of the entire sample. Fluorescence peaks due to ingested chlorophyll are easy to quantify for sample sizes of 20–200 animals. This method obviates some of the problems, such as confinement and previous feeding history, associated with extrapolation from laboratory experiments to field conditions. The method is illustrated by an investigation of diel feeding patterns of marine copepods. Initial results indicate the complexity of these patterns, with either a single night peak or dawn and dusk peaks for adult copepods. Diel variations in feeding activity appear to occur in addition to, but closely interacting with, the diel vertical migrations of the animals.     

Fish predators, food availability and diel vertical migration in Daphnia

Diel vertical migration of zooplankton is a highly variableand complex behaviour which apparently cannot be explained byany single factor. We determined the relative importance offish predation, food availability and water depth in shapingthe migratory behaviour of Daphnia. A modified 2x2x2 factorialexperiment provided two levels of fish density (present/absent),food availability (ambient/high) and depth (4–10 m); shallowtreatments with fish were excluded. Triplicate 1.2 m diameterenclosures for each of the six treatments were held in an 18unit array in Peter Lake, Gogebic Co., MI, USA. Repeated measuresANOVA identified significant trends in daphnid density, migrationand fitness (determined by lipid-ovary-egg index, LOE) as wellas in chlorophyll a content of the water column for part ofa 4-week experiment in July 1988. In deep enclosures with fish,Daphnia performed significantly more intense migrations thanin fishless enclosures, save those in fishless ambient-foodenclosures. Daphnia in deep fishless enclosures without abundantfood at depth performed significantly reduced migrations. DaphnidLOE index was significantly influenced only by food content.Our results were consistent with the predator-avoidance hypothesisas well as with observations of greatest migrations where largevertical differences in food abundance exist. They support ahierarchical view of vertical migration, with presence of fishthe primary factor, and food availability the secondary factor.     

Short-term changes in zooplankton community in Paso Ancho basin (Strait of Magellan): functional trophic structure and diel vertical migration

Knowledge on community structure oriented to describe energy flow during late summer season in Paso Ancho basin (Strait of Magellan) is scarce and particularly affected by vertical diel migration (VDM). The main aim of this work is to identify the VDM patterns of selected species and functional feeding groups of mesozooplankton, collected by the electronic multinet BIONESS in 1995. Detailed studies were carried out on keystone components of the community: 7 species of copepods (Ctenocalanus citer, Drepanopus forcipatus, Metridia lucens, Clausocalanus brevipes, Scolecithricella minor, Paraeuchaeta antarctica, Calanus simillimus), one ostracod (Discoconchoecia elegans), one chaetognath (Sagitta tasmanica), one euphausiid (Euphausia vallentini), and two polychaetes (Pelagobia longicirrata, Tomopteris planktonis). Unexpected deviations from the classic pattern reported in literature were uncovered. The shallow layer mesozooplankton at night, although dominated by exclusively or preferentially herbivores, particularly by E. vallentini, was not represented by several species known as herbivores that remained in the deep layer throughout the day cycle. The deep-layer zooplankton throughout the day cycle was well represented by carnivores, detritivores, and omnivores. It is suggested that during low-chlorophyll summer conditions, the composition of functional groups and diet, and VDM patterns changed to take advantage of sinking phytoplankton and picoplankton in deep layers. Pelagic-benthic coupling would be strengthened due to animals that suppressed their vertical daily rise to the shallow layer at nights but remained in the deep layers to feed on a rain of particulate organic matter and other non-migrant zooplankton.     

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.     

Partial diel vertical migration in an omnivorous macroinvertebrate,Mysis diluviana

Hydrobiologia - Partial migration, whereby only a portion of a population migrates, has just recently received attention in aquatic systems. Partial diel vertical migration (DVM) has received even...     

Concomitant diel vertical migration of a predatory leech and its amphipod prey

SUMMARY.

• 1 The concomitant diet vertical migrations of the predaceous leech Erpobdella montezuma and its pelagic amphipod prey (Hyalella montezuma) were monitored for 1 year in Montezuma Well, Arizona, U.S.A.
• 2 High densities of H. montezuma occurred in the subsurface strata of the water column during the day, but a substantial portion of the population migrated into the surface 1 m at sunset. E. montezuma remained in the lower water strata during the day, but migrated vertically after sunset to exploit the high densities of H. montezuma near the surface. Densities of E. montezuma progressively increased in the upper strata of the water column after sunset as light energy fell below 1 μEin m^?2 s^?1.
• 3 It is suggested that the synchronized nocturnal migrations of the predator E. montezuma in relation to its prey H. montezuma, increases the foraging efficiency of the leech in the highly predictable environment of Montezuma Well.

     

A mechanism of predator-mediated induction of diel vertical migration in Daphnia hyalina

Predator avoidance is generally thought to be the most importantultimate reason for diel vertical migration in zooplankton.Combining field observations and experimental results, it isshown, first, that a phototactic reaction to relative changesin light intensity is at the base of a diel vertical migrationand, secondly, how this phototaxis is enhanced by a chemicalmediated by juvenile perch.     

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.     

Relationships between geotaxis/phototaxis and diel vertical migration in autotrophic dinoflagellates

Marine dinoflagellate diel vertical migrations are often conceptuallyexplained by a species' geotactic and phototactic preferences,but actual simultaneous measurements are rare. Newly collectedsimultaneous measurements on Heterocapsa (Cachonina) illdefina(Herman and Sweeney) and Gymnodinium breve (Davis) are combinedwith similar literature information on Amphidinium carterae(Hulbert), Peridinium faeroense (Paulsen) and Prorocentrum micans(Ehrenberg) to explore several examples of the actual relationshipsbetween diel vertical migration and geotaxis/phototaxis. Amphidiniumcarterae does not migrate, but it exhibits a negative geotaxisthat may counter a small sinking velocity. The four other speciesall exhibit diel vertical migrations that yield surface aggregationsduring daylight, but the associated combinations of geotaxisand phototaxis precision (which is strongest when every cellin a population exhibits the same response to a stimulus andweakest when the response is random) and sign [which is positive(negative) when motion is toward (away from) the stimulus] aredifferent in each case. These different taxis combinations maybe related to species-specific sensor structure and/or placement.Furthermore, variations in the different biochemical pools overa species' cell cycle may contribute to structural/mechanicalchanges that influence how a given sensory array functions ata given time. If so, this coupling may be an important linkin the growth optimization mechanisms and occasional bloom successesof different autotrophic dinoflagellate species under varyingenvironmental conditions.