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Diel vertical migration
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Daily migration of marine life between the twilight zone
and the ocean surface – animation by NASA [1]

Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The adjective "diel" (IPA: /ˈdaɪ.əl/, /ˈdiː.əl/) comes from Latin: diēs, lit. 'day', and refers to a 24-hour period. The migration occurs when organisms move up to the uppermost layer of the water at night and return to the bottom of the daylight zone of the oceans or to the dense, bottom layer of lakes during the day.[2] DVM is important to the functioning of deep-sea food webs and the biologically-driven sequestration of carbon.[3]

In terms of biomass, DVM is the largest synchronous migration in the world.[4][2] It is not restricted to any one taxon, as examples are known from crustaceans (copepods),[5] molluscs (squid),[6] and ray-finned fishes (trout).[7]

The phenomenon may be advantageous for a number of reasons, most typically to access food and to avoid predators.[8] It is triggered by various stimuli, the most prominent being changes in light-intensity,[8] though evidence suggests that biological clocks are an underlying stimulus as well.[9] While this mass migration is generally nocturnal, with the animals ascending from the depths at nightfall and descending at sunrise, the timing can alter in response to the different cues and stimuli that trigger it. Some unusual events impact vertical migration: DVM can be absent during the midnight sun in Arctic regions[10][11] and vertical migration can occur suddenly during a solar eclipse.[12] The phenomenon also demonstrates cloud-driven variations.[13]

The common swift is an exception among birds in that it ascends and descends into high altitudes at dusk and dawn, similar to the vertical migration of aquatic lifeforms.

Discovery

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The phenomenon was first documented by French naturalist Georges Cuvier in 1817. He noted that daphnia, a type of plankton, appeared and disappeared according to a diurnal pattern.[4][14]

During World War II the U.S. Navy was taking sonar readings of the ocean when they discovered the deep scattering layer (DSL). While performing sound propagation experiments, the University of California's Division of War Research (UCDWR) consistently had results of the echo-sounder that showed a distinct reverberation that they attributed to mid-water layer scattering agents. At the time, there was speculation that these readings may be attributed to enemy submarines.[15]

Martin W. Johnson of Scripps Institution of Oceanography proposed a possible explanation. Working with the UCDWR, the Scripps researchers were able to confirm that the observed reverberations from the echo-sounder were in fact related to the diel vertical migration of marine animals. The DSL was caused by large, dense groupings of organisms, like zooplankton, that scattered the sonar to create a false or second bottom.[4][14][15]

Once scientists started to do more research on what was causing the DSL, it was discovered that a large range of organisms were vertically migrating. Most types of plankton and some types of nekton have exhibited some type of vertical migration, although it is not always diel. These migrations may have substantial effects on mesopredators and apex predators by modulating the concentration and accessibility of their prey (e.g., impacts on the foraging behavior of pinnipeds[16]).

Types of vertical migration

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Diel

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This is the most common form of vertical migration. Organisms migrate on a daily basis through different depths in the water column. Migration usually occurs between shallow surface waters of the epipelagic zone and deeper mesopelagic zone of the ocean or hypolimnion zone of lakes.[2] There are three recognized types of diel vertical migration:

Nocturnal vertical migration

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In the most common form, nocturnal vertical migration, organisms ascend to the surface around dusk, remaining at the surface for the night, then migrating to depth again around dawn.[8]

Reverse migration

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Reverse migration occurs with organisms ascending to the surface at sunrise and remaining high in the water column throughout the day until descending with the setting sun.[8]

Twilight diel vertical migration

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Twilight diel vertical migration involves two separate migrations in a single 24-hour period, with the first ascent at dusk followed by a descent at midnight, often known as the "midnight sink". The second ascent to the surface and descent to the depths occurs at sunrise.[8]

Seasonal

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Organisms are found at different depths depending on what season it is.[17] Seasonal changes to the environment may influence changes to migration patterns. Normal diel vertical migration occurs in species of foraminifera throughout the year in the polar regions; however, during the midnight sun, no differential light cues exist so they remain at the surface to feed upon the abundant phytoplankton, or to facilitate photosynthesis by their symbionts.[11] This is not true for all species at all times, however. Zooplankton have been observed to resynchronize their migrations with the light of the moon during periods when the sun is not visible, and to stay in deeper waters when the moon is full.[4]

Larger seasonally-migrating zooplankton such as overwintering copepods have been shown to transport a substantial amount of carbon to the deep ocean through a process known as the lipid pump.[18] The lipid pump is a process that sequesters carbon (in the form of carbon-rich lipids) out of the surface ocean via the descent of copepods to the deep during autumn.[18] These copepods accumulate these lipids during late summer and autumn before descending to the deep to overwinter in response to reduced primary production and harsh conditions at the surface.[18][19] Furthermore, they rely on these lipid reserves that are metabolized for energy to survive through winter before ascending back to the surface in the spring, typically at the onset of a spring bloom.[18]

Ontogenetic

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Organisms spend different stages of their life cycle at different depths.[20] There are often pronounced differences in migration patterns of adult female copepods, like Eurytemora affinis, which stay at depth with only a small upward movement at night, compared to the rest of its life stages which migrate over 10 meters. In addition, there is a trend seen in other copepods, like Acartia spp. that have an increasing amplitude of their DVM seen with their progressive life stages. This is possibly due to increasing body size of the copepods and the associated risk of visual predators, like fish, as being larger makes them more noticeable.[5]

Vertical migration stimuli

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There are two different types of factors that are known to play a role in vertical migration, endogenous and exogenous. Endogenous factors originate from the organism itself; sex, age, size, biological rhythms, etc. Exogenous factors are environmental factors acting on the organism such as light, gravity, oxygen, temperature, predator-prey interactions, etc.[21]

Endogenous factors

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Endogenous rhythm

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Biological clocks are an ancient and adaptive sense of time innate to an organism that allows them to anticipate environmental changes and cycles so they are able to physiologically and behaviorally respond to the expected change.[9]

Evidence of circadian rhythms controlling DVM, metabolism, and even gene expression have been found in copepod species, Calanus finmarchicus. These copepods were shown to continue to exhibit these daily rhythms of vertical migration in the laboratory setting even in constant darkness, after being captured from an actively migrating wild population.[9]

An experiment was done at the Scripps Institution of Oceanography which kept organisms in column tanks with light/dark cycles. A few days later the light was changed to a constant low light and the organisms still displayed diel vertical migration. This suggests that some type of internal response was causing the migration.[22]

Clock gene expression

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Many organisms, including the copepod C. finmarchicus, have genetic material devoted to maintaining their biological clock. The expression of these genes varies temporally with the expression significantly increasing following dawn and dusk at times of greatest vertical migration. These findings may indicate they work as a molecular stimulus for vertical migration.[9]

Body size

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The relative body size of an organism has been found to affect DVM. Bull trout express daily and seasonal vertical migrations with smaller individuals always staying at a deeper layer than the larger individuals. This is most likely due to a predation risk, but is dependent on the individuals own size such that smaller animals may be more inclined to remain at depth.[7]

Exogenous factors

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Light

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"Light is the most common and critical cue for vertical migration".[8] However, as of 2010, there had not been sufficient research to determine which aspect of the light field was responsible.[8] As of 2020, research has suggested that both light intensity and spectral composition of light are important.[23]

Temperature

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Organisms will migrate to a water depth with temperatures that best suit the organisms needs, for example some fish species migrate to warmer surface waters in order to aid digestion. Temperature changes can influence swimming behavior of some copepods. In the presence of a strong thermocline some zooplankton may be inclined to pass through it, and migrate to the surface waters, though this can be very variable even in a single species. The marine copepod, Calanus finmarchicus, will migrate through gradients with temperature differences of 6 °C over George's Bank; whereas, in the North Sea they are observed to remain below the gradient.[24]

Salinity

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Changes in salinity may promote organism to seek out more suitable waters if they happen to be stenohaline or unequipped to handle regulating their osmotic pressure. Areas that are impacted by tidal cycles accompanied by salinity changes, estuaries for example, may see vertical migration in some species of zooplankton.[25] Salinity has also been proposed as a factor that regulates the biogeochemical impact of diel vertical migration.[26]

Pressure

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Pressure changes have been found to produce differential responses that result in vertical migration. Many zooplankton will react to increased pressure with positive phototaxis, a negative geotaxis, and/or a kinetic response that results in ascending in the water column. Likewise, when there is a decrease in pressure, the zoo plankton respond by passively sinking or active downward swimming to descend in the water column.[25]

Predator kairomones

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A predator might release a chemical cue which could cause its prey to vertically migrate away.[27] This may stimulate the prey to vertically migrate to avoid said predator. The introduction of a potential predator species, like a fish, to the habitat of diel vertical migrating zooplankton has been shown to influence the distribution patterns seen in their migration. For example, a study used Daphnia and a fish that was too small to prey on them (Lebistus reticulatus), found that with the introduction of the fish to the system the Daphnia remained below the thermocline, where the fish was not present. This demonstrates the effects of kairomones on Daphnia DVM.[24]

Tidal patterns

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Some organisms have been found to move with the tidal cycle. A study looked at the abundance of a species of small shrimp, Acetes sibogae, and found that they tended to move further higher in the water column and in higher numbers during flood tides than during ebb tides experiences at the mouth of an estuary. It is possible that varying factors with the tides may be the true trigger for the migration rather than the movement of the water itself, like the salinity or minute pressure changes.[25]

Reasons for vertical migration

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There are many hypotheses as to why organisms would vertically migrate, and several may be valid at any given time.[28]

Predator avoidance

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The universality of DVM suggests that there is some powerful common factor behind it. The connection between available light and DVM has led researchers to theorize that organisms may stay in deeper, darker areas during the day to avoid being eaten by predators who depend on light to see and catch their prey. While the ocean's surface provides an abundance of food, it may be safest for many species to visit it at night.[4]

Light-dependent predation by fish is a common pressure that causes DVM behavior in zooplankton and krill. A given body of water may be viewed as a risk gradient whereby the surface layers are riskier to reside in during the day than deep water, and as such promotes varied longevity among zooplankton that settle at different daytime depths.[29] Indeed, in many instances it is advantageous for zooplankton to migrate to deep waters during the day to avoid predation and come up to the surface at night to feed. For example, the northern krill Meganyctiphanes norvegica undergoes diel vertical migration to avoid planktivorous fish.[30]

Patterns among migrators seem to support the predator avoidance theory. Migrators will stay in groups as they migrate, a behavior that may protect individuals within the group from being eaten. Groups of smaller, harder to see animals begin their upward migration before larger, easier to see species, consistent with the idea that detectability by visual predators is a key issue. Small creatures may start to migrate upwards as much as 20 minutes before the sun sets, while large conspicuous fish may wait as long as 80 minutes after the sun goes down. Species that are better able to avoid predators also tend to migrate before those with poorer swimming capabilities. Squid are a primary prey for Risso's dolphins (Grampus griseus), an air-breathing predator, but one that relies on acoustic rather than visual information to hunt. Squid delay their migration pattern by about 40 minutes when dolphins are about, lessening risk by feeding later and for a shorter time.[4][31]

Metabolic advantages

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Another possibility is that predators can benefit from diel vertical migration as an energy conservation strategy. Studies indicate that male dogfish (Scyliorhinus canicula) follow a "hunt warm - rest cool" strategy that enables them to lower their daily energy costs. They remain in warm water only long enough to obtain food, and then return to cooler areas where their metabolism can operate more slowly.[31][32][33]

Alternatively, organisms feeding on the bottom in cold water during the day may migrate to surface waters at night in order to digest their meal at warmer temperatures.[34]

Dispersal and transport

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Organisms can use deep and shallow currents to find food patches or to maintain a geographical location.

Avoid UV damage

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The sunlight can penetrate into the water column. If an organism, especially something small like a microbe, is too close to the surface the UV can damage them. So they would want to avoid getting too close to the surface, especially during daylight.[35][36]

Water transparency

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A theory known as the "transparency-regulator hypothesis" predicts that "the relative roles of UV and visual predation pressure will vary systematically across a gradient of lake transparency."[35][36] In less transparent waters, where fish are present and more food is available, fish tend to be the main driver of DVM. In more transparent bodies of water, where fish are less numerous and food quality improves in deeper waters, UV light can travel farther, thus functioning as the main driver of DVM in such cases.[37]

Unusual events

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Due to the particular types of stimuli and cues used to initiate vertical migration, anomalies can change the pattern drastically.

For example, the occurrence of midnight sun in the Arctic induces changes to planktonic life that would normally perform DVM with a 24-hour night and day cycle. In the summers of the Arctic the Earth's north pole is directed toward the sun creating longer days and at the high latitude continuous day light for more than 24-hours.[10] Species of foraminifera found in the ocean have been observed to cease their DVM pattern, and rather remain at the surface in favor of feeding on the phytoplankton.[38] For example Neogloboquadrina pachyderma, and for those species that contain symbionts, like Turborotalita quinqueloba, remain in sunlight to aid photosynthesis.[11] Changes in sea-ice and surface chlorophyll concentration are found to be stronger determinants of the vertical habitat of Arctic N. pachyderma.[38]

There is also evidence of changes to vertical migration patterns during solar eclipse events. In the moments that the sun is obscured during normal day light hours, there is a sudden dramatic decrease in light intensity. The decreased light intensity, replicates the typical lighting experienced at night time that stimulate the planktonic organisms to migrate. During an eclipse, some copepod species distribution is concentrated near the surface, for example Calanus finmarchicus displays a classic diurnal migration pattern but on a much shorter time scale during an eclipse.[12]

Importance for the biological pump

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Biological pump
Diel vertically migrating krill, smaller zooplankton such as copepods, and fish can actively transport carbon to depth by consuming particulate organic carbon (POC) in the surface layer at night, and metabolising it at their daytime, mesopelagic residence depths. Depending on species life history, active transport may occur on a seasonal basis as well.[39]

The biological pump is the conversion of CO2 and inorganic nutrients by plant photosynthesis into particulate organic matter in the euphotic zone and transference to the deeper ocean.[40] This is a major process in the ocean and without vertical migration it wouldn't be nearly as efficient. The deep ocean gets most of its nutrients from the higher water column when they sink down in the form of marine snow. This is made up of dead or dying animals and microbes, fecal matter, sand and other inorganic material.

Organisms migrate up to feed at night so when they migrate back to depth during the day they defecate large sinking fecal pellets.[40] Whilst some larger fecal pellets can sink quite fast, the speed that organisms move back to depth is still faster. At night organisms are in the top 100 metres of the water column, but during the day they move down to between 800 and 1000 meters. If organisms were to defecate at the surface it would take the fecal pellets days to reach the depth that they reach in a matter of hours. Therefore, by releasing fecal pellets at depth they have almost 1000 metres less to travel to get to the deep ocean. This is known as active transport. The organisms are playing a more active role in moving organic matter down to depths. Because a large majority of the deep sea, especially marine microbes, depends on nutrients falling down, the quicker they can reach the ocean floor the better.

Zooplankton and salps play a large role in the active transport of fecal pellets. 15–50% of zooplankton biomass is estimated to migrate, accounting for the transport of 5–45% of particulate organic nitrogen to depth.[40] Salps are large gelatinous plankton that can vertically migrate 800 meters and eat large amounts of food at the surface. They have a very long gut retention time, so fecal pellets usually are released at maximum depth. Salps are also known for having some of the largest fecal pellets. Because of this they have a very fast sinking rate, small detritus particles are known to aggregate on them. This makes them sink that much faster. As previously mentioned, the lipid pump represents a substantial flux of POC (particulate organic carbon) to the deep ocean in the form of lipids produced by large overwintering copepods.[18] Through overwintering, these lipids are transported to the deep in autumn and are metabolized at depths below the thermocline through winter before the copepods rise to the surface in the spring.[18] The metabolism of these lipids reduces this POC at depth while producing CO2 as a waste product, ultimately serving as a potentially significant contributor to oceanic carbon sequestration.[18] Although the flux of lipid carbon from the lipid pump has been reported to be comparable to the global POC flux from the biological pump, observational challenges with the lipid pump from deficient nutrient cycling,[41][42][43] and capture techniques have made it difficult to incorporate it into the global carbon export flux.[19][44][45] So while currently there is still much research being done on why organisms vertically migrate, it is clear that vertical migration plays a large role in the active transport of dissolved organic matter to depth.[46]

See also

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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia

Diel vertical migration

View on Grokipedia
from Grokipedia
Diel vertical migration (DVM), also known as diurnal vertical migration, is a synchronized daily movement pattern observed in aquatic environments, where planktonic and nektonic organisms—such as , copepods, euphausiids, , and cephalopods—ascend to surface waters at to feed and descend to deeper layers before dawn, often covering distances from hundreds of meters to over 800 meters in the . This behavior is ubiquitous across marine and freshwater systems, from epipelagic to bathypelagic zones, and represents the largest migration on , involving billions of individuals globally. The migration is primarily driven by a balance between opportunities and predation risk, with organisms exploiting nutrient-rich surface layers under the cover of darkness while retreating to deeper, darker depths during daylight to evade visually predators. Light intensity serves as the key environmental cue, often aligning with specific isolumes (e.g., around 10⁻³ W m⁻²), though internal circadian rhythms, gradients, prey availability, and oxygen levels can modulate the timing and amplitude of movements. In some cases, such as with certain dinoflagellates or deep-sea , migrations may also optimize or reduce exposure to harmful radiation. Ecologically, DVM profoundly influences aquatic food webs and biogeochemical cycles, comprising 40–60% of mesozooplankton in many regions and forming detectable deep-scattering layers via . By transporting organic carbon from surface productivity to the through active and fecal pellets, it enhances the , contributing 15–40% to particle export and up to 50% of respiration in migration layers, thereby playing a critical role in global and nutrient recycling. This process also generates , affects , and supports higher trophic levels, underscoring DVM's integral role in maintaining dynamics and influencing regulation.

Introduction

Definition and scope

Diel vertical migration (DVM) refers to the synchronized daily vertical displacement of aquatic organisms, in which they typically ascend from deeper waters to surface layers at dusk and descend back to depth , following a 24-hour cycle tightly linked to the light-dark transition. This behavior represents a fundamental adaptive strategy in aquatic ecosystems, enabling organisms to optimize and amid varying environmental conditions. The scope of DVM encompasses primarily planktonic and nektonic organisms, including zooplankton such as copepods and , micronekton like small fish and mysids, and various larval stages. It is distinct from random or irregular vertical movements, as DVM exhibits precise, rhythmic patterns driven by predictable environmental cycles rather than factors. This phenomenon is observed across marine and freshwater systems, though it is most pronounced in oceanic environments where light penetration structures the . Basic patterns of DVM include migration amplitudes ranging from 100 to 800 meters in oceanic settings, with organisms often phase-locked to the —ascending shortly before sunset and descending before sunrise to align with optimal conditions. The behavior is highly ubiquitous, affecting the vast majority of species and, in some systems, a substantial portion of the biomass participates in these migrations. In terms of ecological scale, DVM constitutes one of the largest migrations on , involving an estimated ~5 billion tons of organisms and dwarfing all terrestrial animal migrations combined.

Global occurrence and key examples

Diel vertical migration (DVM) is a ubiquitous phenomenon observed across diverse aquatic ecosystems worldwide, encompassing marine, freshwater, and estuarine environments, and involving a broad array of organisms from to . In marine settings, it is particularly prominent in the open ocean, where vast populations of and micronekton undertake synchronized daily movements spanning hundreds of meters. For instance, (Euphausia superba) in the perform extensive DVM, descending to depths of 200–400 meters during the day and ascending to surface waters at night, facilitating nutrient transport and predator avoidance in these polar regions. Similarly, mesopelagic fishes such as (family Myctophidae) dominate the deep scattering layers of the world's oceans, migrating from depths of 500–750 meters by day to the upper 200 meters at night, contributing significantly to the global biomass flux in the . In freshwater systems, DVM is equally widespread, especially among cladocerans in temperate lakes, where it helps regulate predator-prey dynamics in stratified water columns. A key example is the water flea Daphnia species, which commonly migrate vertically by 5–20 meters daily, residing in deeper, cooler hypolimnetic layers during daylight to evade visual predators like fish and rising to the epilimnion at night for foraging on phytoplankton. This behavior is documented in numerous lakes across North America and Europe, underscoring its role in maintaining ecosystem balance in these enclosed basins and driving vertical transport of organic matter at ecosystem scales. Estuarine and coastal habitats exhibit more variable DVM patterns, often modified by tidal influences and salinity gradients in shallow bays. Mysids (Crustacea: Mysidae), such as Mysis diluviana, demonstrate partial DVM in these dynamic environments, with only a fraction of the population migrating from benthic substrates to the water column at night, while others remain in place, allowing niche partitioning and adaptation to fluctuating conditions. This partial migration is observed in coastal systems like those around the and North American estuaries, where it supports energy transfer between seafloor and surface layers. The scale of DVM is immense, representing one of the planet's largest synchronized animal movements by biomass, with global oceanic fluxes estimated to rival terrestrial migrations in magnitude and involving trillions of individuals nightly. Recent studies up to 2025 highlight ongoing observations of DVM in changing environments; for example, in subarctic epipelagic waters off , capelin (Mallotus villosus) and cod (Gadus morhua) exhibit pronounced diel movements that structure feeding interactions within the . Likewise, larval and juvenile small yellow croaker (Larimichthys polyactis) in coastal waters of the River Estuary, , perform DVM to optimize transport and survival, ascending at night to exploit faster currents for offshore dispersal. These examples illustrate the adaptability and ecological significance of DVM across global aquatic realms.

Historical discovery

Early observations

The earliest documented observations of diel vertical migration emerged in the , primarily through the work of naturalists employing rudimentary sampling techniques in both freshwater and marine environments. In 1817, French naturalist provided the first description of the phenomenon in the freshwater cladoceran , noting its retreat to deeper waters at midday to avoid intense light and its ascent toward the surface in the evening as light diminished. This observation highlighted a rhythmic, light-driven , though it was initially confined to shallow lakes and lacked broader ecological context. Subsequent marine explorations built on such anecdotal records, shifting focus to oceanic plankton. Marine records began with depth-stratified net sampling during mid-century expeditions. In the 1850s, British naturalist Edward Forbes, during his expeditions in the aboard vessels like HMS Beacon, used dredges and tow nets to collect and benthic organisms, observing progressive changes in species abundance and composition with depth, emphasizing zonation patterns where surface-dwelling appeared scarcer during daylight hours compared to nighttime hauls, though he attributed variations partly to tidal influences rather than strictly diel cycles. These qualitative insights marked an initial recognition of vertical structuring in Mediterranean communities, influencing later oceanographic surveys. Systematic evidence from global voyages further illuminated the phenomenon. The HMS Challenger expedition (1872–1876), a landmark British scientific circumnavigation, conducted repeated tows at varying depths and times across the Atlantic, Pacific, and Indian Oceans, documenting diurnal variations in catches from deep-sea hauls—such as increased surface abundance of crustaceans and radiolarians at night. Reports by expedition naturalists, including John Murray, noted these patterns in open-ocean settings, linking them to light availability and providing the first global-scale hints of widespread diel migration among pelagic organisms. Concurrently, late-19th-century fisheries observations offered practical corroboration; for instance, 1880s observations from Scottish fisheries, such as those by Brook (1886) in , noted daytime near-surface aggregations of copepods like that decreased with presence, suggesting behavioral responses to predation exploited by fishermen but initially unexplained beyond seasonal cues. Despite these pioneering efforts, early methods imposed significant limitations on understanding. Reliance on manual net tows—often coarse-meshed and deployed sporadically—yielded inconsistent data, capturing only discrete depth intervals and missing the continuous, real-time dynamics of migration. Such techniques, as used by and Challenger scientists, frequently confounded diel patterns with tidal or weather effects, resulting in incomplete profiles until the advent of more precise, continuous sampling in the . These foundational observations nonetheless established diel vertical migration as a recurrent ecological feature, later validated through acoustic and optical technologies.

Scientific advancements and key researchers

In the early 20th century, significant methodological progress in studying diel vertical migration (DVM) came from the development of quantitative sampling tools, notably the Clarke-Bumpus plankton sampler introduced in the 1930s by George L. Clarke. This device allowed for precise, depth-specific collections of plankton, enabling researchers to document vertical distributions and migration patterns with greater accuracy than previous net hauls, thus providing foundational quantitative data on DVM amplitudes in oceanic waters. Clarke, a pioneering oceanographer, advanced quantitative sampling and hypotheses on light-driven migrations during , such as the rate-of-change hypothesis linking DVM to irradiance variations, marking a shift from descriptive to mechanistic observations of subsurface behaviors. A major breakthrough occurred during when sonar operators detected moving 'false bottoms' in the ocean, later identified as deep scattering layers (DSLs) formed by migrating and , providing the first acoustic evidence of widespread DVM. Building on this, the mid-20th century saw John E. Harris's influential work in the 1950s and 1960s, where he demonstrated the role of endogenous rhythms in driving DVM through laboratory experiments on copepods, isolating circadian clocks from external cues like light. Harris's 1963 paper established that internal biological timers persist under constant conditions, solidifying the endogenous component of migration control. By the 1960s, electronic depth sounders enhanced resolution of migration dynamics, revealing daily vertical excursions of up to several hundred meters in layers, which previous sampling had underestimated. The advent of acoustic Doppler current profilers (ADCPs) in the 2000s revolutionized real-time tracking, allowing non-invasive monitoring of DVM across large water columns by detecting acoustic from migrating organisms. Milestone expeditions like the Tara Oceans voyage (2009–2013) integrated multidisciplinary sampling to map global DVM patterns, combining net tows, acoustics, and imaging to quantify vertical fluxes and community structures in the upper ocean. Recent modeling efforts in the 2020s, such as the 2025 study by Vilain et al., which used Continuous Plankton Recorder data and Map Equation clustering to show how DVM influences bioregions in the North Atlantic, linking behavioral patterns to large-scale biogeographic boundaries. Technological evolution has progressed from discrete net-based sampling to advanced satellite telemetry, which tracks individual migrators' depths via pop-up tags, and genomic sequencing, revealing genes associated with light sensitivity and circadian regulation in DVM performers. These innovations, exemplified by high-throughput eDNA analysis, now enable identification of migration-related genetic adaptations across taxa.

Types of vertical migration

Diel patterns

Diel vertical migration primarily manifests in three distinct patterns among aquatic organisms, particularly : nocturnal, reverse, and twilight migrations. Nocturnal migration represents the standard and most prevalent subtype, in which organisms ascend from deeper waters to the surface layers around dusk, remain in the upper throughout the night, and descend to deeper depths by dawn. This behavior is exhibited by the majority of zooplankton species in marine environments, including many copepods and euphausiids. For instance, nearly all mesopelagic zooplankton engage in this pattern, contributing to massive daily displacements across basins. Reverse migration, a less common variant, involves organisms rising to shallower depths during the day and sinking to deeper layers at night. This subtype is observed in specific taxa, such as juveniles or males in certain species, and in some freshwater cladocerans. Examples include smaller individuals of Pseudocalanus spp. in coastal bays, which follow this inverted cycle relative to the dominant nocturnal pattern. Twilight migration features limited vertical excursions synchronized with dawn and dusk transitions, typically involving partial ascents and descents confined to shallow depths with amplitudes under 50 m. This pattern predominates in tidally influenced coastal regions, such as among mysid crustaceans like Mysis spp., where movements remain relatively shallow and do not span the full . Overall, nocturnal migration dominates oceanic systems, while reverse and twilight forms are more regionally specific, with the latter often tied to dynamic nearshore habitats. These diel patterns may exhibit seasonal variations in amplitude or timing.

Non-diel patterns

Non-diel vertical migrations encompass patterns that operate on timescales longer than the daily cycle, including seasonal and ontogenetic shifts in depth distribution among . Seasonal vertical migration (SVM) involves gradual depth changes over months, often driven by annual environmental variations, where organisms descend to deeper waters during unfavorable periods such as winter for overwintering. For instance, large polar copepods like and C. glacialis exhibit SVM with amplitudes reaching over 1000 m in polar seas, relocating from surface layers in summer to depths exceeding 500 m in winter to conserve energy in . Ontogenetic vertical migration (OVM) refers to progressive depth shifts across an individual's lifespan, correlating with developmental stages and body size increases. In calanoid copepods such as Eucalanus inermis, naupliar stages remain near the surface to access , while copepodite and stages descend deeper, often into oxygen minimum zones below 200 m, with deepening continuing through maturation. This pattern is prevalent in many long-lived species, including approximately half of calanoid copepod taxa, where OVM facilitates stage-specific resource utilization and predator avoidance. These non-diel patterns interact with diel vertical migration by establishing baseline depths that modulate daily amplitudes; for example, seasonal overwintering descent in polar copepods reduces the scope for diel excursions during winter, while deepening in juveniles limits their participation in full adult diel cycles. Body size effects, linked to , further influence these shifts by affecting metabolic demands and capacity. Such interactions highlight how extended migrations provide a framework for daily behaviors in diverse aquatic ecosystems.

Controls on diel migration

Endogenous mechanisms

Endogenous mechanisms underlying diel vertical migration (DVM) primarily involve internal biological clocks that drive and synchronize migratory behaviors, independent of immediate environmental fluctuations. These circadian rhythms, with free-running periods typically ranging from 23 to 25 hours, enable organisms to anticipate daily cycles and initiate vertical movements even in constant conditions. In the freshwater cladoceran species, for instance, these rhythms persist under constant darkness, maintaining synchronized ascent and descent patterns that mirror natural DVM. Similarly, in marine copepods like , laboratory experiments demonstrate endogenous oscillations in swimming activity and respiration that continue for several days in the absence of light-dark cues, confirming the role of an internal timing mechanism. At the molecular level, these rhythms are governed by genes, such as period (per) and (cry), which form transcriptional-translational feedback loops in s. These loops oscillate to regulate physiological processes, including adjustments and behaviors critical for DVM. In (Euphausia superba), daily peaks in per and cry expression align with migratory phases, driving metabolic and locomotor changes that facilitate vertical positioning. Orthologs of these genes, conserved across arthropods, have been identified in the of , underscoring their ancient role in synchronizing behaviors. Body size influences the expression of these endogenous mechanisms through metabolic scaling, where larger individuals often exhibit greater migration amplitudes or deeper depths due to allometric relationships. For example, larger copepods descend to greater depths during daylight, as observed in field studies. This arises from endogenous programming that adjusts vertical excursions to optimize individual fitness. Supporting evidence for these internal drivers comes from controlled studies and genetic manipulations. Experiments in perpetual darkness reveal persistence, as seen in ostracods (Asterope and Philomedes) where circadian swimming patterns endure without external zeitgebers. Furthermore, genetic knockouts of clock s disrupt DVM timing; a 2024 study in Daphnia showed that per gene inactivation abolishes sustained vertical migration under constant conditions, highlighting the clock's essential role in generation. These findings collectively affirm that endogenous clocks provide a robust, heritable framework for DVM across taxa.

Exogenous cues

Light serves as the primary exogenous cue, or , regulating the timing of diel vertical migration in most aquatic organisms, with changes in intensity and spectral quality triggering ascent and descent phases. , increasing blue light wavelengths (approximately 450-500 nm) prompt descent to deeper waters, enabling organisms to avoid visually hunting predators during daylight hours, while , red-shifted facilitates ascent to surface layers for feeding. This response is highly sensitive, with migration often initiating at low intensities around 10^{-7} μmol photons m^{-2} s^{-1} for species such as Calanus and , with thresholds varying from 10^{-8} to 10^{-6} μmol photons m^{-2} s^{-1} across taxa and habitats. Temperature gradients, particularly thermal stratification in the , modulate the speed and extent of migration, often accelerating movements in response to warmer surface layers. In stratified environments, organisms like copepods exhibit faster ascent rates in epilimnetic waters (typically 5-15°C warmer than deeper layers), which can enhance overall migration efficiency by reducing exposure time in vulnerable zones. For instance, in species such as , warmer temperatures increase swimming velocities, allowing quicker transitions between depths. Additional exogenous factors include gradients in estuarine systems, which guide orientation and vertical positioning during migration. In coastal zones with sharp haloclines, lower in surface waters can cue upward movements, helping larvae and juveniles maintain position against tidal flows. Some crustaceans sense hydrostatic pressure changes, providing depth feedback that fine-tunes descent depths and prevents overshooting. Chemical cues from predators, known as kairomones, further enhance descent behaviors, particularly in freshwater and coastal like , where these infochemicals induce deeper positioning even under low light conditions to evade detection. In coastal areas, tidal cycles synchronize migrations, with tides often aligning with ascent to exploit nutrient-rich inflows. Interactions among these cues emphasize light's dominance, yet can amplify its effects; for example, thermal gradients of 5-10°C across the have been observed to roughly double migration rates in dinoflagellates and copepods by boosting metabolic and responses to signals. Such synergies ensure adaptive precision in dynamic aquatic environments.

Adaptive functions

Diel vertical migration (DVM) primarily functions to minimize encounters with visually oriented predators, such as planktivorous , by exploiting variations in availability across depths. During daylight hours, migrating and descend to depths exceeding 200 meters, where penetration is limited and visual foraging by predators is severely impaired, thereby reducing predation risk in the well-lit surface layers. At night, ascent to shallower waters occurs under conditions of low ambient , further limiting the effectiveness of sight-based predation. This temporal and spatial separation is a key adaptive strategy observed across diverse aquatic systems, from freshwater lakes to oceanic environments. In addition to light-mediated behaviors, chemical signals known as —released by predators like —play a crucial role in enhancing DVM as a predator avoidance mechanism, particularly in species such as . These infochemicals induce rapid and intensified migratory responses, prompting individuals to increase the depth or amplitude of their descent during the day, which strengthens separation from predators. For instance, the bile salt 5α-cyprinol , a specific kairomone from , elicits significant daytime downward shifts in at concentrations as low as 100 pM, amplifying the behavioral response beyond light cues alone. This chemical induction allows prey to fine-tune their migration based on perceived predation threat, often resulting in more pronounced avoidance patterns. While DVM provides clear anti-predator benefits, it incurs metabolic costs associated with active swimming, which can elevate respiration rates and consume a substantial portion of the daily energy budget in some . These costs are offset by substantial survival advantages; modeling studies indicate that DVM can substantially reduce predatory mortality when visual predators dominate, far outweighing the energetic expenditure. Field and experiments further demonstrate that non-migrating individuals experience markedly higher predation rates than migrants, due to prolonged exposure in vulnerable surface waters. Reverse DVM patterns, where prey ascend during the day and descend at night, also emerge in systems dominated by non-visual predators, such as chaetognaths, to minimize overlap with these nocturnal hunters and protect vulnerable life stages.

Physiological and foraging advantages

Diel vertical migration provides significant physiological benefits to migrating organisms, particularly through metabolic optimization enabled by thermal stratification in aquatic environments. By ascending to warmer surface waters at night for feeding and descending to cooler deeper layers during the day, zooplankton such as copepods and daphnids experience reduced respiration rates in the colder depths, leveraging the Q10 effect where metabolic rates typically decrease by a factor of 2–3 for every 10°C drop in temperature. For instance, at 10°C compared to 20°C, metabolic rates can be 2–3 times lower, allowing energy that would otherwise be expended on maintenance to be allocated toward growth and reproduction. This strategy can yield significant daily energy savings relative to constant surface residency, assuming equal time spent in warm and cold layers and a Q10 of 2, as modeled in early bioenergetic analyses. Foraging advantages further enhance the adaptive value of diel migration, as nighttime surface excursions grant access to concentrated and microzooplankton resources that are scarce in deeper waters. In many systems, is confined to the euphotic zone, enabling migrators like calanoid copepods to ingest substantially more carbon during brief surface sojourns compared to what would be available at depth. This pulsed feeding optimizes energy intake while minimizing time in risky surface conditions, supporting higher growth and reproductive output over non-migratory conspecifics. Migration also mitigates physiological stress from (UV) radiation, a potent DNA-damaging agent in sunlit surface layers. Daytime descent to depths below 10 m in clear waters reduces UVB exposure by approximately 99%, as attenuation in even oligotrophic conditions limits penetrable UV to shallow euphotic zones. Some species, including daphnids, further adapt through inducible pigmentation changes, such as increased production, to shield against residual UV during ascent or in shallower migrations. In clear oligotrophic waters, where visual challenges arise due to high transparency, diel migration enhances feeding efficiency by timing surface access to nocturnal low-light periods, when reduced visibility aids prey capture without compromising the benefits of deeper refuge. This transparency-driven pattern allows migrators to exploit sparse surface resources more effectively than in turbid systems, where such vertical shifts are less pronounced.

Ecological roles

Carbon cycling and

Diel vertical migration (DVM) contributes to the ocean's through active transport of carbon, where migrating organisms, primarily and micronekton, feed on and particulate organic carbon in surface waters at night before descending to deeper layers during the day, where they respire , excrete , and produce fast-sinking fecal pellets. This process decouples from remineralization, effectively exporting carbon from the euphotic zone to the mesopelagic, with active flux via DVM estimated to account for 15–20% of passive particulate organic carbon in many regions. Globally, DVM-mediated carbon is quantified at approximately 0.5–1.3 Gt C year⁻¹, primarily through respiration, fecal pellet production, and excretion, representing about 10% of total oceanic carbon . Recent modeling of micronekton DVM, using trait-based approaches that account for size-dependent production rates, indicates these contribute 10–18% to total active relative to passive sinking, with fishes showing the highest efficiency due to deeper migrations and rapid gut processing, particularly in temperate regions during summer. DVM amplifies the biological pump by enhancing overall carbon sequestration; one-dimensional models demonstrate that migration-induced decoupling of feeding and remineralization increases deep export by approximately 15% compared to non-migratory scenarios, with twilight zones (100–200 m depths) acting as hotspots where up to 50% of total respiration occurs due to concentrated migrant biomass. This active component supplements passive particle sinking, elevating net flux to the deep ocean and supporting long-term carbon storage. Empirical evidence from sediment trap deployments correlates DVM amplitude with elevated carbon export, as traps capture increased fecal pellet fluxes during periods of high migration intensity, often exceeding passive sinking rates. In the North Atlantic, studies on copepod DVM, particularly of , reveal seasonal lipid storage and migrations export 2–6 g C m⁻² year⁻¹ via , comparable to sinking fluxes at 600–1,400 m depths and verified against trap data showing repackaged in pellets.

Food web dynamics and biogeography

Diel vertical migration (DVM) plays a pivotal role in structuring aquatic food webs by facilitating the transfer of energy and nutrients between surface and deeper layers, effectively linking epipelagic primary production to mesopelagic consumers. Migrating zooplankton and micronekton, such as euphausiids, ascend to the surface at night to feed on phytoplankton and then descend during the day, exporting organic matter downward through respiration, excretion, and predation mortality. This process supports higher trophic levels in the mesopelagic zone, where non-migratory fishes and invertebrates rely on the influx of surface-derived biomass. For instance, Antarctic krill (Euphausia superba), which exhibit pronounced DVM, serve as a primary prey for baleen whales like humpback and fin whales, channeling epipelagic energy to these large predators and sustaining their populations across polar food webs. DVM also influences dispersal and transport mechanisms within aquatic systems, promoting vertical and horizontal of organisms via interactions with currents. During ascent and descent, migrants can be passively transported by layered currents, enhancing connectivity between distant habitats. Recent studies on larval small yellow croaker () in the demonstrate how DVM allows juveniles to exploit faster currents at specific depths, facilitating offshore transport and success. Similarly, in the North Atlantic, communities exhibit DVM-driven separation timescales that define distinct bioregions, with day-night depth partitioning altering species distributions and connectivity patterns. On a broader biogeographic scale, DVM contributes to the delineation of bioregions through depth-based partitioning, creating four primary zones in regions like the North Atlantic where migratory behaviors synchronize community structures diurnally. This partitioning influences gradients, as variations in migration amplitude and timing along latitudinal or depth clines modulate coexistence and resource use. Furthermore, DVM affects success by imposing barriers to range expansion; for example, tropical vertical migrators face challenges crossing cold-water thermal gradients during migrations, limiting poleward invasions and shaping global distribution patterns. The ecological impacts of DVM extend to fisheries and interspecies interactions, altering predation dynamics and niche overlap. In the Barents Sea, capelin (Mallotus villosus) DVM influences cod (Gadus morhua) predation efficiency, as capelin's daytime descent into deeper waters reduces overlap with surface-oriented cod schools, affecting stock assessments and fishery yields. Partial migrations, where only subsets of populations migrate, further enable niche partitioning; in mysids (Mysis spp.), stable isotope analysis reveals that migrating individuals exploit pelagic resources while non-migrators access benthic food sources, reducing intraspecific competition and enhancing overall community stability.

Variations and anomalies

Environmental influences on patterns

Diel vertical migration (DVM) patterns vary significantly across aquatic habitats due to differences in water column depth and physical dynamics. In freshwater lakes, migration amplitudes are typically shallower, ranging from 10 to 50 meters, constrained by the limited depth and stronger thermal stratification compared to oceanic environments where amplitudes can extend from 200 to 1000 meters or more, allowing for broader vertical excursions in response to and predation gradients. In estuarine systems, tidal cycles introduce additional synchronization that often reduces the regularity of diel patterns, as and larval migrations align more closely with tidal flows for retention or transport, overriding pure solar-driven rhythms. Climate change exerts profound influences on DVM through ocean warming and acidification, altering migration amplitudes and cue perception. Warming waters may reduce migration amplitudes in zooplankton due to changes in temperature tolerance, as observed in freshwater systems where elevated temperatures decrease migratory range. Ocean acidification causes copepods to avoid low-pH water, potentially restricting them to shallower areas and affecting predator avoidance behaviors. Anthropogenic pollution, particularly artificial light at night, further modifies DVM timing in coastal and urban-adjacent waters. Light pollution from urban sources can alter the timing and amplitude of migrations in taxa like by interfering with natural photoperiod signals. Recent observations highlight ongoing environmental shifts in DVM patterns. In regions, (Mallotus villosus) primarily occupy 0–60 m depths both day and night, with a small fraction of juveniles exhibiting reverse DVM. Similarly, mysid demonstrate partial DVM for niche partitioning, where subsets of populations remain in benthic or pelagic zones day and night, influenced by gradients and resource competition in lakes like Superior.

Unusual migrations in non-aquatic systems

While diel vertical migration (DVM) is predominantly studied in aquatic environments, analogous behaviors have been observed in non-aquatic systems, particularly among microbes and terrestrial . In alpine snowpacks, motile microbes such as and exhibit DVM over distances of 10-20 cm daily, driven by phototaxis to avoid intense daytime solar radiation and diurnal meltwater flows that position them near the surface (0-3 cm) at night and deeper layers (10-20 cm) during the day. A 2025 study in a northern Japanese alpine forest documented this pattern in (e.g., Chloromonas sp.) and associated , where surface positioning at night supports nutrient access in melt layers and enhances photosynthetic activity under reduced light. Terrestrial soil , such as collembolans (springtails), display diel shifts in vertical distribution within soil profiles, though these are not true DVM but rather responses to surface conditions. In arable soils, Arthropleona collembolans like Lepidocyrtus cyaneus show peak abundances at the soil surface from midday to midnight, correlated with higher soil-surface temperatures, with reduced surface activity (likely deeper positioning) during cooler night hours, minimizing exposure risks. In aquatic contexts, unusual DVM variants occur beyond typical zooplankton, including reverse patterns in fish. A 2025 investigation in a subarctic Greenland epipelagic ecosystem revealed that juvenile capelin (Mallotus villosus) exhibit reverse DVM, with smaller individuals (<10 cm) migrating from shallow depths (0-60 m) at night to deeper layers (80-120 m) during the day, potentially to evade cod (Gadus morhua) predation while foraging on copepods; this contrasts with the standard ascent of larger capelin. In hypoxic zones, DVM amplitudes reach extremes, as seen in the Red Sea's mesopelagic layers where scattering layers of fish and invertebrates migrate ~700 m vertically—from 400-600 m daytime depths through oxygen minima (<1.4 mL L⁻¹) to <200 m at night—tolerating low oxygen via physiological adaptations like high hemoglobin affinity. Disrupted DVM cues can lead to anomalous events, such as increased stranding risks for nearshore organisms. Artificial suppresses standard DVM in , causing avoidance of illuminated surface layers and reduced upward migration at night, potentially heightening vulnerability to predation. Additionally, a 2025 modeling study in the system quantified separation timescales for vertically migrating , showing DVM facilitates faster divergence from passive particles over seasonal timescales, potentially altering local distributions under variable currents.

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