Marine snow

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, which is referred to as the biological pump. Export production is the amount of organic matter produced in the ocean by primary production that is not recycled (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. mg C m−2 d−1). The term was coined by explorer William Beebe as observed 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 photosynthetic activity and ocean currents. Marine snow can be an important food source for organisms living in the aphotic zone, particularly for organisms that 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 phytoplankton, protists, fecal matter, sand, and other inorganic dust. Most trapped particles are more vulnerable to grazers than they would be as free-floating individuals. Aggregates can form through abiotic processes (i.e. extrapolymeric substances). These are natural polymers exuded as waste products mostly by phytoplankton and bacteria. Mucus secreted by zooplankton (mostly salps, appendicularians, and pteropods) also contributes to the constituents of marine snow aggregates. 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 blooms. As phytoplankton 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. Most organic components of marine snow are consumed by microbes, 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 ecosystems: 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). 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.

Aggregate dynamics Aggregates begin as the colloidal fraction, which typically contains particles sized between one nanometer 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 bioavailable.

Ballasting effect

Aggregates that sink more quickly to the bottom of the ocean have a greater chance of exporting carbon to the deep sea floor. The longer the residence time in the water column the greater the chance of being grazed upon. Aggregates formed in high dust areas are able to increase their densities faster and in more superficial layers compared to aggregates formed without dust particles present and these aggregates with increased lithogenic material have also been correlated with particulate organic carbon fluxes, however when they become heavily ballasted with lithogenic material they cannot scavenge any additional minerals during their descent, which suggests that carbon export to the deep ocean in regions with high dust deposition is strongly controlled by dust input to the surface ocean while suspended dust particles in deeper water layers do not significantly interact with sinking aggregates.

Fragmentation 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. During zooplankton's vertical migration, the abundances of aggregates increased while size distributions decreased. Aggregates were found in the abdomen in zooplankton indicating their grazing will fragment larger aggregates.

Surface coagulation Aggregates may also form from colloids trapped on the surface of rising bubbles. For example, Kepkay et al. found that bubble coagulation leads to an increase in bacterial respiration since more food is available to them.

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.

Particle-associated microorganisms

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