Diel vertical migration

Diel vertical migration, also known as diurnal vertical migration, is a pattern of movement that some organisms living in the ocean and in lakes undertake each day. Usually organisms move up to the epipelagic zone at night and return to the mesopelagic zone of the oceans or to the hypolimnion zone of lakes during the day. The word diel comes from the Latin dies day, and means a 24-hour period.

Discovery

During World War II the Navy was taking sonar readings of the ocean when they discovered the deep scattering layer (DSL). The DSL was caused by large groupings of organisms that scattered the sonar to create a false or second bottom. The false bottom was shallower during the day and deeper during the night; this was the first recording of diel vertical migration.

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 (fish) have exhibited some type of vertical migration, although it is not always diel.

Types and stimuli of vertical migration

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- An experiment was done at the Scripps Oceanographic Research Institute 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. Thus suggestions that some type of internal response was causing the migration.^[1]

Exogenous factors^[2]

• Light - An organisms wants 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 or during an eclipse they will start to migrate.
• Temperature - sometimes thermoclines can act as a barrier that an organism will not cross.
• Salinity - in areas such as the Arctic melting ice was cause a layer of freshwater which organisms cannot cross.
• Predator kairomones - a predator might release a chemical cue which could cause its prey to vertically migrate away.^[3]

Types of vertical migration

• Diel - This has already been mentioned and is the most common form. Organisms migrate daily, usually up to shallow waters at night and deep waters during the day. But there are other forms of vertical migration worth mentioning.
• Seasonal - Organisms are found at different depths depending on what season it is.^[4]
• Ontogentic - Organisms spend different stages of their life cycle at different depths.^[5]

Reasons to vertically migrate

There are a lot of hypotheses on why organisms would vertically migrate. Each of these hypotheses are equally valid and most of the time more than one of the hypothesis account for a reason a species would vertical migrate.^[6]

• Predator avoidance - Organisms come up to shallow waters at night to feed while it’s dark out because their predators cannot see them as easily.
• 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 actually damage them. So they would want to avoid getting too close to the surface, especially during daylight.

Importance with 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 transferred to the deeper ocean.^[7] 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 higher up in the water column and it sinks down in the form of marine snow. Marine snow is made up of dead or dying animals and microbes, fecal matter, sand, and other inorganic material. Fecal matter is what vertical migration helps get to depth much faster.

Organisms migrate up to feed at night so when they migrate back to depth during the day they defecate large sinking fecal pellets.^[7] 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 800–1000 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 are to play a large role in the active transport of fecal pellets. 15-50% of zooplankton biomass above 15-50% is estimated to vertically migrate and this can account for the transport of 5-45% of Particulate Organic Nitrogen to depth.^[7] As for salps, they are large gelatinous plankton that can vertically migrate 800 meters and eat large amounts of food at the surface. They also 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 and small detritus particles are known to aggregate on them. This makes them sink that much faster. So while currently there is still a lot of research being done on why organisms vertically migrate, it is very clear that vertical migration plays a large role in the active transport of dissolved organic matter to depth.^[8]

See also

• Krill
• Phytoplankton
• Plankton
• Primary production

Reference

1. ^ Enright, J.T.; W.M. Hammer (1967). "Vertical Diurnal Migration and Endogenous Rhythmicity". Science 157 (3791): 937–941. doi:10.1126/science.157.3791.937. JSTOR 1722121. PMID 17792830. 
2. ^ Richards, Shane; Hugh Possingham, John Noye (1996). "Diel vertical migration: modeling light-mediated mechanisms". Journal of Plankton Research 18 (12): 2199–2222. doi:10.1093/plankt/18.12.2199. http://plankt.oxfordjournals.org/content/18/12/2199.full.pdf+html. 
3. ^ von Elert, Eric; Georg Pohnert (2000). "Diel Predator specificity of kairomones in diel vertical migration of Daphnia: a chemical approach". OIKOS 88 (1): 119–128. doi:10.1034/j.1600-0706.2000.880114.x. ISSN 0030-1299. http://0-www.jstor.org.helin.uri.edu/sici?sici=0030-1299%282000%2988%3A1%3C119%3APSOKID%3E2.0.CO%3B2-X&origin=serialsolutions. 
4. ^ Visser, Andre; Sigrun Jonasdottir (1999). "Lipids, buoyancy and the seasonal vertical migration of Calanus finmarchicus". Fisheries Oceanography 8: 100–106. doi:10.1046/j.1365-2419.1999.00001.x. 
5. ^ Kobari, Toru; Tsutomu Ikeda (2001). "Octogenetic vertical migration and life cycle of Neocalanus plumchrus (Crustacea:Copepoda) in the Oyashio region, with notes on regional variations in body size". Journal of Plankton Research 23 (3): 287–302. doi:10.1093/plankt/23.3.287. http://plankt.oxfordjournals.org/content/23/3/287.full.pdf+html. 
6. ^ Kerfoot, WC (1985). "Adaptive value of vertical migration: Comments on the predation hypothesis and some alternatives". Contributions in Marine Science 27: 91–113. http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=1587950. 
7. ^ ^{a b c} Steinberg, Deborah; Sarah Goldthwait, Dennis Hansell (2002). "Zooplankton vertical migration and the active transport of dissolved organic and inorganic nitrogen in the Sargasso Sea". Deep-Sea Research I 49 (8): 1445–1461. doi:10.1016/S0967-0637(02)00037-7. ISBN 1493597039. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6VGB-46HFXF2-5&_user=657938&_coverDate=08%2F31%2F2002&_rdoc=1&_fmt=high&_orig=search&_origin=search&_sort=d&_docanchor=&view=c&_searchStrId=1493597039&_rerunOrigin=google&_acct=C000035679&_version=1&_urlVersion=0&_userid=657938&md5=cce92453ae63045d33db450ec07edbf1&searchtype=a. 
8. ^ Wiebe, P.H; L.P. Madin, L.R. Haury, G.R. Harbison, L.M. Philbin (1979). "Diel Vertical Migration by Salpa aspera and its potential for large-scale particulate organic matter transport to the deep-sea". Marine Biology 53 (3): 249–255. doi:10.1007/BF00952433. http://www.springerlink.com/content/j25h1553175j2174/.