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Diel vertical migration

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 migration occurs
when organisms move up to the
uppermost layer of the sea 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. The word diel comes from
the Latin dies day, and means a 24-
hour period. In terms of biomass, it
is the greatest migration in the Daily migration of marine life to and from the twilight zone
world.[2] It is not restricted to any to the ocean surface – animation by NASA [1]
one taxon as examples are known
from crustaceans (copepods),[3]
molluscs (squid),[4] and ray-finned fishes (trout).[5] Various stimuli are responsible for this
phenomenon, the most prominent being response to changes in light intensity,[6] though evidence
suggests that biological clocks are an underlying stimulus as well.[7] The phenomenon may arise for a
number of reasons, though it is most typically to access food and avoid predators.[6] While this mass
migration is generally nocturnal, with the animals ascending from the depths at nightfall and descending
at sunrise, the timing can be altered in response to the different cues and stimuli that trigger it. Some
unusual events impact vertical migration: DVM is absent during the midnight sun in Arctic regions[8]
and vertical migration can occur suddenly during a solar eclipse.[9]

Contents
Discovery
Types of vertical migration
Vertical migration stimuli
Endogenous factors
Endogenous rhythm
Clock gene expression
Body size
Exogenous factors
Light
Temperature
Salinity
Pressure
Predator kairomones
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Tidal patterns
Reasons for vertical migration
Unusual events
Importance for the biological pump
See also
References

Discovery
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. By collaborating with
biologists from Scripps Institution and the UCDWR, they 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.[10]

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[11]).

Types of vertical migration

Diel
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]:
Furthermore, there are three recognized types of diel vertical migration. This includes nocturnal
vertical migration, the most common form, where organisms ascend to the surface around dusk,
remaining at the surface for the night, then migrating to depth again around dawn. The second
form is reverse migration, which 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. The
third form is twilight diel vertical migration, involving 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.[6]
Seasonal
Organisms are found at different depths depending on what season it is.[12] 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.[8]
Ontogenetic
Organisms spend different stages of their life cycle at different depths.[13] There are often
pronounced differences in migration patterns of adult female copepods, like Eurytemora affinis,
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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.[3]

Vertical migration stimuli

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

Endogenous factors

Endogenous rhythm

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. 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.[7] 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.[14]

Clock gene expression

Many organisms including the copepod C. finmarchicus, has genetic material devoted to maintaining its
biological clock. The expression of these genes varies temporally with the expression significantly
increasing following dawn and dusk at times of greatest vertical migration seen in this species. These
findings may indicate they work as a molecular stimulus for vertical migration.[7]

Body size

The relative body size of an organism has been found to effect 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.[5]

Exogenous factors

Light

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Light is the most common and critical cue for vertical migration.[6] Organisms want to find an optimum
light intensity (isolume). Whether it is no light or a large amount of light, an organism will travel to
where it is most comfortable. Studies have shown that during a full moon organisms will not migrate up
as far, and that during an eclipse they will start to migrate.[15]

Temperature

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.[16]

Salinity

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.[17] In areas such as the Arctic, melting ice causes a layer of freshwater which
organisms cannot cross.

Pressure

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.[17]

Predator kairomones

A predator might release a chemical cue which could cause its prey to vertically migrate away.[18] 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 of 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.[16]

Tidal patterns

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.[17]
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Reasons for vertical migration

There are many hypotheses as to why organisms would vertically migrate, and several may be valid at
any given time.[19]

Predator avoidance
Light-dependent predation by fish is a common pressure that causes DVM behavior in
zooplankton. 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.[20] 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.
Metabolic advantages
By feeding in the warm surface waters at night and residing in the cooler deep waters during the
day they can conserve energy. 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.
Dispersal and transport
Organisms can use deep and shallow currents to find food patches or to maintain a geographical
location.
Avoid UV damage
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.

Water transparency

A recent theory of DVM, termed the Transparency Regulator Hypothesis, argues that water transparency
is the ultimate variable that determines the exogenous factor (or combination of factors) that causes
DVM behavior in a given environment.[21] 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.[22]

Unusual events
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. Species of foraminifera found in the ocean cease their DVM pattern, and rather
remain at the surface in favor of feeding on the phytoplankton, for example Neogloboquadrina
pachyderma, and for those species that contain symbionts, like Turborotalita quinqueloba, remain in
sunlight to aid photosynthesis.[8]

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

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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.[9]

Importance for the biological pump

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.[24] 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 Biological pump
large sinking fecal pellets.[24] Whilst Diel vertically migrating krill, smaller zooplankton such as copepods, and
some larger fecal pellets can sink fish can actively transport carbon to depth by consuming particulate
organic carbon (POC) in the surface layer at night, and metabolising it at
quite fast, the speed that organisms
their daytime, mesopelagic residence depths. Depending on species life
move back to depth is still faster. At
history, active transport may occur on a seasonal basis as well.[23]
night organisms are in the top 100
metres of the water column, but
during the day they move down to
between 800–1000 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 something 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.[24] 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. 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.[25]

See also
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Krill
Phytoplankton
Primary production

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