View source for Arrival theorem

<!-- Deleted image removed: [[Image:RLampitt Pelagra recovery.jpg|thumb|right|300px|[[Underway]] recovery of a [[sediment trap]] used to collect marine snow and estimate export production]] -->
In the deep ocean, '''marine snow''' is a continuous shower of mostly organic [[detritus]] falling from the upper layers of the [[water column]]. It is a significant means of exporting [[energy]] from the [[light]]-rich [[photic zone]] to the [[aphotic zone]] below.  The term was first coined by the explorer [[William Beebe]] as he observed it from his [[bathysphere]].  As the origin of marine snow lies in activities within the productive photic zone, the prevalence of marine snow changes with seasonal fluctuations in [[photosynthesis|photosynthetic]] activity and [[ocean current]]s. Marine snow can be an important food source for organisms living in the aphotic zone, particularly for organisms which live very deep in the water column.

==Composition==
Marine snow is made up of a variety of mostly organic matter, including dead or dying animals and plants ([[plankton]]), [[protist]]s ([[diatom]]s), fecal matter, sand, soot and other inorganic dust. Most trapped particles are more vulnerable to grazers than they would be as free floating individuals and can be classified as "olive green" or "gray body" cells, which are plant parts and degrading plant material.  A majority of marine snow composition is actually made up of aggregates of smaller particles held together by a sugary mucus, transparent extracellular polysaccharides (TEPs).  These are natural [[polymer]]s exuded as waste products mostly by [[phytoplankton]] and [[bacterium|bacteria]]. [[Mucus]] secreted by [[zooplankton]] (mostly [[salps]], [[appendicularia]]ns, and [[pteropoda|pteropods]]) also contribute to the constituents of marine snow aggregates.<ref>{{cite book | last=Miller | first=Charles B. | title=Biological Oceanography | year=2004 | publisher=Blackwell Science Ltd.| pages=94–5, 266–7}}</ref>  These aggregates grow over time and may reach several centimeters in diameter, traveling for weeks before reaching the ocean floor.

Marine snow often forms during [[algal bloom]]s.  As algae accumulate, they aggregate or get captured in other aggregates, both of which accelerate the sinking rate.  Aggregation and sinking is actually thought to be a large component of sources for algae loss from surface water.<ref>{{cite book | last=Mann| first=K.H. |last=Lazier |first=J.R.N | title=Dynamics of Marine Ecosystems | year=2006 | publisher=Blackwell Publishing| page=35}}</ref>  Most organic components of marine snow are consumed by [[microbe]]s, [[zooplankton]] and other filter-feeding animals within the first 1,000 metres of their journey. In this way marine snow may be considered the foundation of deep-sea [[mesopelagic]] and [[benthic]] [[ecosystem]]s: As sunlight cannot reach them, deep-sea organisms rely heavily on marine snow as an energy source. The small percentage of material not consumed in shallower waters becomes incorporated into the muddy "ooze" blanketing the ocean floor, where it is further decomposed through biological activity.

Marine snow aggregates exhibit characteristics that fit Goldman's "aggregate spinning wheel hypothesis".  This hypothesis states that phytoplankton, microorganisms and bacteria live attached to aggregate surfaces and are involved in rapid nutrient recycling.  Phytoplankton have been shown to be able to take up nutrients from small local concentrations of organic material (e.g. fecal matter from an individual zooplankton cell, regenerated nutrients from organic decomposition by bacteria).<ref>{{cite journal | author=McCarthy J.J, Goldman J.C. | title=Nitrogenous nutrition of marine phytoplankton in nutrient-depleted waters | doi=10.1126/science.203.4381.670  | year=1979 | journal=[[Science (journal)|Science]] | volume=203 | pages=670–672 }}</ref>  As the aggregates slowly sink to the bottom of the ocean, the many microorganisms residing on them are constantly respiring and contribute greatly to the [[microbial loop]].

==Aggregation==
Aggregates begin as the colloidal fraction, which typically contains particles sized between 1&nbsp;nm and several micrometers.  The colloidal fraction of the ocean contains a large amount of organic matter unavailable to grazers.  This fraction has a much higher total mass than either phytoplankton or bacteria but is not readily available due to size characteristics of the particles in relation to potential consumers.  The colloidal fraction must aggregate in order to be more [[bioavailability|bioavailable]].  Aggregation theory outlines main mechanisms by which marine aggregates may form and are as follows:
:[[Brownian motion]]
::Brownian motion, also known as the Brownian Pump in this case, describes the interaction between individual particles in the colloid fraction.  Particles in Brownian motion interact randomly due to the impact of solute molecules.  These interactions lead to collision and aggregation of these small particles.  The small aggregates then collide and aggregate with other aggregates and particles until the aggregate in question is several micrometers in diameter.
:[[shear (fluid)|Shear]]
::Once particles have aggregated to several micrometers in diameter, they begin to accumulate bacteria, since there is sufficient site space for feeding and reproduction.  At this size it is large enough to undergo sinking.  It also has the components necessary to fit the "aggregate spinning wheel hypothesis." Evidence for this has been found by Alldredge and Cohen (1987) who found evidence of both respiration and photosynthesis within aggregates, suggesting the presence of both [[autotrophic]] and [[heterotrophic]] organisms.<ref>{{cite journal | author=Alldredge, A.L. and Cohen, Y. | title=Can microscale chemical patches persist in the sea?  Microelectrode study of marine snow, fecal pellets. | year=1987 | journal=Science | volume=235| pages=689–691 | doi = 10.1126/science.235.4789.689 | issue=4789}}</ref>
:Differential [[Settling]]
::This form of aggregation involves particles sinking at different rates and their collision to form aggregates.
:[[diffusion|Diffusive]] Capture
::Diffusive capture describes a particle that is advected into the diffusion-limited boundary layer of another particle and is eventually captured/collides with that particle.
:Surface [[Coagulation]]
::Aggregates may also form from colloids trapped on the surface of rising [[Liquid bubble|bubble]]s.  For example, Kepkay et al. found that bubble coagulation leads to an increase in bacterial respiration since more food is available to them.<ref>{{cite journal | author= Kepkay, P.E.| title=Particle aggregation and the biological activity of colloids| year=1994 | journal=Marine Ecology Project Series | volume=109 | pages=293–304 }}</ref>  In fact, a storm at sea could increase bacterial respiration by up to 36 fold for two to four hours.
:[[Filtration]]
::Particles and small organisms floating through the water column can become trapped within aggregates.  Marine snow aggregates are porous, however, and some particles are able to pass through them.
:Bacterial Motility
::It is unclear how relative this mode of aggregation is to marine snow, but there have been limited observations of bacteria moving fast and far enough to capture colloidal particles.

Aggregation theory represents a two state system.  At low cell concentrations aggregation is relatively unimportant and somewhat more unlikely.  However, at higher cell concentrations it becomes increasingly important.  A model has been proposed to characterize the formation of marine aggregates and the loss due to sinking:

:<math>\frac{\mathrm{d} C_{1} }{\mathrm{d} t} = g C_1 - 20.8\alpha r^{3} G C_1^2|</math>

:''C<sub>1</sub>'' is the concentration of the cells
:''r'' is the radius of each cell
:''G'' is the shearing rate
:''&alpha;'' is the stickiness coefficient
:''g'' is the growth rate.

Thus, aggregation of marine particles is more prevalent when cell and particle concentration is higher (e.g. algal blooms)

==Study==
'''Export production''' is the amount of [[organic matter]] produced in the [[ocean]] by [[primary production]] that is not recycled ([[remineralisation|remineralised]]) before it sinks into the [[aphotic zone]].  Because of the role of export production in the ocean's [[biological pump]], it is typically measured in units of [[carbon]] (e.g. [[milligram|mg]] C [[square metre|m<sup>-2</sup>]] [[day|d<sup>-1</sup>]]).

The fraction of primary production that is exported to the aphotic zone is generally higher when primary production occurs in short (seasonal) bursts, than when it occurs more evenly spread out across a year.

Because of the relatively long residence time of the ocean's [[thermohaline circulation]], carbon transported as marine snow into the deep ocean by the [[biological pump]] can remain out of contact with the atmosphere for more than 1000 years.  That is, when the marine snow is finally decomposed to inorganic nutrients and dissolved [[carbon dioxide]], these are effectively isolated from the surface ocean for relatively long time-scales related to [[ocean circulation]].  Consequently, enhancing the quantity of marine snow that reaches the deep ocean is the basis of several [[geoengineering]] schemes to enhance [[carbon sequestration]] by the ocean.  [[Ocean nourishment]] and [[iron fertilisation]] seek to boost the production of organic material in the surface ocean, with a concomitant rise in marine snow reaching the deep ocean.<ref>{{cite journal | last=Lampitt | first=Richard | coauthors=''et al.'' | title=Ocean fertilisation: a potential means of geoengineering? | year=2008 | journal=[[Philosophical Transactions of the Royal Society|Phil. Trans. Roy. Soc. A]] | volume=366 | pages=3919–3945 | doi=10.1098/rsta.2008.0139 }}</ref>  These efforts have not yet produced a sustainable fertilization that effectively transports carbon out of the system.

Increases in ocean temperatures, a projected indicator of [[climate change]], may result in a decrease in the production of marine snow due to the enhanced [[stratification (water)|stratification]] of the water column.  Increasing stratification decreases the availability of phytoplankton nutrients such as [[nitrate]], [[phosphate]] and [[silicic acid]], and could lead to a decrease in primary production and, thus, marine snow.

Marine snow has also begun to garner interest from [[microbiology|microbiologists]], owing to the microbial communities associated with it. Recent research indicates transported bacteria may exchange [[gene]]s with previously thought to be isolated populations of bacteria inhabiting the breadth of the ocean floor. In such an immense area there may be as yet undiscovered species tolerant of high pressures and extreme cold, perhaps finding use in [[bioengineering]] and [[pharmacy]].

== See also ==
* [[Biological pump]]
* [[Detritivore]]
* [[f-ratio]]
* [[Marine mucilage]]
* [[Sediment trap]]
* [[Whale fall]]

==References==
{{reflist}}

== External links ==
* [http://www.spaceref.com/news/viewpr.html?pid=5897 SpaceRef.com, Deep sea bacteria get new genes from marine snow]
* [http://halodule.marsci.uga.edu/Research/marine_snow.html U. Georgia, Marine Snow and Particles]
* [http://www.sos.bangor.ac.uk/marine/mb/O3B14/Marine_snow.html U. Bangor, Marine Snow: Formation and composition]
* [http://www.niwa.cri.nz/pubs/wa/09-4/potential NIWA, What grows up must fall down: the potential impact of climate change on plankton and carbon export]
* [http://lepo.it.da.ut.ee/~olli/eutr/html/htmlBook_63.html Primary production and vertical export]

{{aquatic ecosystem topics|expanded=marine}}
{{physical oceanography|expanded=none}}
{{fishery science topics}}

[[Category:Aquatic ecology]]
[[Category:Bioindicators]] <!-- a bioindicator of what exactly? -->
[[Category:Biological oceanography]]
[[Category:Chemical oceanography]]
[[Category:Ecological processes]]
[[Category:Fisheries science]]
[[Category:Geochemistry]]
[[Category:Oceanography]]