plankton

Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish and whales.

Marine plankton include bacteria, archaea, algae, protozoa and drifting or floating animals that inhabit the saltwater of oceans and the brackish waters of estuaries. Freshwater plankton are similar to marine plankton, but are found in the freshwaters of lakes and rivers. Plankton are usually thought of as inhabiting water, but there are also airborne versions, the aeroplankton, that live part of their lives drifting in the atmosphere. These include plant spores, pollen and wind-scattered seeds, as well as microorganisms swept into the air from terrestrial dust storms and oceanic plankton swept into the air by sea spray.

Though many planktonic species are microscopic in size, ''plankton'' includes organisms over a wide range of sizes, including large organisms such as jellyfish. Plankton are defined by their ecological niche and level of motility rather than by any phylogenetic or taxonomic classification. Technically the term does not include organisms on the surface of the water, which are called ''neuston''—or those that swim actively in the water, which are called '' nekton''.

Terminology

The name ''plankton'' is derived from the Greek adjective (), meaning ''errant'', and by extension, ''wanderer'' or ''drifter'', and was coined by Victor Hensen in 1887. While some forms are capable of independent movement and can swim hundreds of meters vertically in a single day (a behavior called diel vertical migration), their horizontal position is primarily determined by the surrounding water movement, and plankton typically flow with ocean currents. This is in contrast to nekton organisms, such as fish, squid and marine mammals, which can swim against the ambient flow and control their position in the environment.

Within the plankton, holoplankton spend their entire life cycle as plankton (e.g. most algae, copepods, salps, and some jellyfish). By contrast, meroplankton are only planktic for part of their lives (usually the larval stage), and then graduate to either a nektic (swimming) or benthic (sea floor) existence. Examples of meroplankton include the larvae of sea urchins, starfish, crustaceans, marine worms, and most fish.

The amount and distribution of plankton depends on available nutrients, the state of water and a large amount of other plankton.

The study of plankton is termed planktology and a planktonic individual is referred to as a plankter. The adjective ''planktonic'' is widely used in both the scientific and popular literature, and is a generally accepted term. However, from the standpoint of prescriptive grammar, the less-commonly used ''planktic'' is more strictly the correct adjective. When deriving English words from their Greek or Latin roots, the gender-specific ending (in this case, "-on" which indicates the word is neuter) is normally dropped, using only the root of the word in the derivation.

Trophic groups

Plankton are primarily divided into broad functional (or trophic level) groups:
* Phytoplankton (from Greek ''phyton'', or plant), are autotrophic prokaryotic or eukaryotic algae that live near the water surface where there is sufficient light to support photosynthesis. Among the more important groups are the diatoms, cyanobacteria, dinoflagellates and coccolithophores.
*Zooplankton (from Greek ''zoon'', or animal), are small protozoans or metazoans (e.g. crustaceans and other animals) that feed on other plankton. Some of the eggs and larvae of larger nektonic animals, such as fish, crustaceans, and annelids, are included here.
*Mycoplankton include fungi and fungus-like organisms, which, like bacterioplankton, are also significant in remineralisation and nutrient cycling.
*Bacterioplankton include bacteria and archaea, which play an important role in remineralising organic material down the water column (note that prokaryotic phytoplankton are also bacterioplankton).
* Virioplankton are viruses. Viruses are more abundant in the plankton than bacteria and archaea, though much smaller.

Mixoplankton

*Mixotrophs. Plankton have traditionally been categorized as producer, consumer and recycler groups, but some plankton are able to benefit from more than just one trophic level. In this mixed trophic strategy—known as mixotrophy—organisms act as both producers and consumers, either at the same time or switching between modes of nutrition in response to ambient conditions. This makes it possible to use photosynthesis for growth when nutrients and light are abundant, but switching to eat phytoplankton, zooplankton or each other when growing conditions are poor. Mixotrophs are divided into two groups; constitutive mixotrophs, CMs, which are able to perform photosynthesis on their own, and non-constitutive mixotrophs, NCMs, which use phagocytosis to engulf phototrophic prey that are either kept alive inside the host cell which benefit from its photosynthesis, or they digest their prey except for the plastids which continues to perform photosynthesis (kleptoplasty).
Recognition of the importance of mixotrophy as an ecological strategy is increasing, as well as the wider role this may play in marine biogeochemistry. Studies have shown that mixotrophs are much more important for the marine ecology than previously assumed, and comprise more than half of all microscopic plankton. Their presence act as a buffer that prevents the collapse of ecosystems during times with little to no light.

Size groups

Plankton are also often described in terms of size. Usually the following divisions are used:
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However, some of these terms may be used with very different boundaries, especially on the larger end. The existence and importance of nano- and even smaller plankton was only discovered during the 1980s, but they are thought to make up the largest proportion of all plankton in number and diversity.

The microplankton and smaller groups are microorganisms and operate at low Reynolds numbers, where the viscosity of water is more important than its mass or inertia.

File:Plankton size.png,

Habitat groups

Marine plankton

Marine plankton includes marine bacteria and archaea, algae, protozoa and drifting or floating animals that inhabit the saltwater of oceans and the brackish waters of estuaries.

Freshwater plankton

Freshwater plankton are similar to marine plankton, but are found inland in the freshwaters of lakes and rivers.

Aeroplankton

File:Ocean mist and spray 2.jpg, Sea spray containing marine microorganisms can be swept high into the atmosphere and may travel the globe as aeroplankton before falling back to earth.

Aeroplankton are tiny lifeforms that float and drift in the air, carried by the current of the wind; they are the atmospheric analogue to oceanic plankton. Most of the living things that make up aeroplankton are very small to microscopic in size, and many can be difficult to identify because of their tiny size. Scientists can collect them for study in traps and sweep nets from aircraft, kites or balloons. Aeroplankton is made up of numerous microbes, including viruses, about 1000 different species of bacteria, around 40,000 varieties of fungi, and hundreds of species of protists, algae, mosses and liverworts that live some part of their life cycle as aeroplankton, often as spores, pollen, and wind-scattered seeds. Additionally, peripatetic microorganisms are swept into the air from terrestrial dust storms, and an even larger amount of airborne marine microorganisms are propelled high into the atmosphere in sea spray. Aeroplankton deposits hundreds of millions of airborne viruses and tens of millions of bacteria every day on every square meter around the planet.

The sea surface microlayer, compared to the sub-surface waters, contains elevated concentration of bacteria and viruses.Blanchard, D.C., 1983. The production, distribution and bacterial enrichment of the sea-salt aerosol. In: Liss, P.S., Slinn, W.G.N. ŽEds.., Air–Sea Exchange of Gases and Particles. D. Reidel Publishing Co., Dordrecht, Netherlands, pp. 407-444. These materials can be transferred from the sea-surface to the atmosphere in the form of wind-generated aqueous aerosols due to their high vapour tension and a process known as volatilisation.Wallace Jr., G.T., Duce, R.A., 1978. Transport of particulate organic matter by bubbles in marine waters. Limnol. Oceanogr. 23 Ž6., 1155–1167. When airborne, these microbes can be transported long distances to coastal regions. If they hit land they can have affect animal, vegetation and human health.WHO, 1998. Draft guidelines for safe recreational water environments: coastal and fresh waters, draft for consultation. World Health Organization, Geneva, EOSrDRAFTr98 14, pp. 207–299. Marine aerosols that contain viruses can travel hundreds of kilometers from their source and remain in liquid form as long as the humidity is high enough (over 70%).Klassen, R. D., & Roberge, P. R. (1999). Aerosol transport modeling as an aid to understanding atmospheric corrosivity patterns. Materials & Design, 20, 159–168.Moorthy, K. K., Satheesh, S. K., & Krishna Murthy, B.V. (1998). Characteristics ofspectral optical depths and size distributions of aerosols over tropical oceanic regions. Journal of Atmospheric and Solar–Terrestrial Physics, 60, 981–992.
These aerosols are able to remain suspended in the atmosphere for about 31 days.Aller, J., Kuznetsova, M., Jahns, C., Kemp, P. (2005) The sea surface microlayer as a source of viral and bacterial enrichment in marine aerosols. Journal of aerosol science. Vol. 36, pp. 801-812. Evidence suggests that bacteria can remain viable after being transported inland through aerosols. Some reached as far as 200 meters at 30 meters above sea level. The process which transfers this material to the atmosphere causes further enrichment in both bacteria and viruses in comparison to either the SML or sub-surface waters (up to three orders of magnitude in some locations).Marks, R., Kruczalak, K., Jankowska, K., & Michalska, M. (2001). Bacteria and fungi in air over the GulfofGdansk and Baltic sea. Journal of Aerosol Science, 32, 237–250.

Geoplankton

Many animals live in terrestrial environments by thriving in transient often microscopic bodies of water and moisture, these include rotifers and gastrotrichs which lay resilient eggs capable of surviving years in dry environments, and some of which can go dormant themselves. Nematodes are usually microscopic with this lifestyle. Water bears, despite only having lifespans of a few months, famously can enter suspended animation during dry or hostile conditions and survive for decades. This allows them to be ubiquitous in terrestrial environments despite needing water to grow and reproduce. Many microscopic crustacean groups like copepods and amphipods (of which sandhoppers are members) and seed shrimp are known to go dormant when dry and live in transient bodies of water too

Other groups

Gelatinous zooplankton

Gelatinous zooplankton are fragile animals that live in the water column in the ocean. Their delicate bodies have no hard parts and are easily damaged or destroyed. Gelatinous zooplankton are often transparent. All jellyfish are gelatinous zooplankton, but not all gelatinous zooplankton are jellyfish. The most commonly encountered organisms include ctenophores, medusae, salps, and Chaetognatha in coastal waters. However, almost all marine phyla, including Annelida, Mollusca and Arthropoda, contain gelatinous species, but many of those odd species live in the open ocean and the deep sea and are less available to the casual ocean observer.

Ichthyoplankton

Ichthyoplankton are the eggs and larvae of fish. They are mostly found in the sunlit zone of the water column, less than 200 metres deep, which is sometimes called the epipelagic or photic zone. Ichthyoplankton are planktonic, meaning they cannot swim effectively under their own power, but must drift with the ocean currents. Fish eggs cannot swim at all, and are unambiguously planktonic. Early stage larvae swim poorly, but later stage larvae swim better and cease to be planktonic as they grow into juveniles. Fish larvae are part of the zooplankton that eat smaller plankton, while fish eggs carry their own food supply. Both eggs and larvae are themselves eaten by larger animals.What are Ichthyoplankton?
Southwest Fisheries Science Center, NOAA. Modified 3 September 2007. Retrieved 22 July 2011. Fish can produce high numbers of eggs which are often released into the open water column. Fish eggs typically have a diameter of about . The newly hatched young of oviparous fish are called larvae. They are usually poorly formed, carry a large yolk sac (for nourishment) and are very different in appearance from juvenile and adult specimens. The larval period in oviparous fish is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on zooplankton prey, a process which depends on typically inadequate zooplankton density, starving many larvae. In time fish larvae become able to swim against currents, at which point they cease to be plankton and become juvenile fish.

Holoplankton

Holoplankton are organisms that are planktic for their entire life cycle. Holoplankton can be contrasted with meroplankton, which are planktic organisms that spend part of their life cycle in the benthic zone. Examples of holoplankton include some diatoms, radiolarians, some dinoflagellates, foraminifera, amphipods, krill, copepods, and salps, as well as some gastropod mollusk species. Holoplankton dwell in the pelagic zone as opposed to the benthic zone. Holoplankton include both phytoplankton and zooplankton and vary in size. The most common plankton are protists.

Meroplankton

Meroplankton are a wide variety of aquatic organisms which have both planktonic and benthic stages in their life cycles. Much of the meroplankton consists of larval stages of larger organism. Meroplankton can be contrasted with holoplankton, which are planktonic organisms that stay in the pelagic zone as plankton throughout their entire life cycle. After a period of time in the plankton, many meroplankton graduate to the nekton or adopt a benthic (often sessile) lifestyle on the seafloor. The larval stages of benthic invertebrates make up a significant proportion of planktonic communities. The planktonic larval stage is particularly crucial to many benthic invertebrate in order to disperse their young. Depending on the particular species and the environmental conditions, larval or juvenile-stage meroplankton may remain in the pelagic zone for durations ranging from hour to months.

Pseudoplankton

Pseudoplankton are organisms that attach themselves to planktonic organisms or other floating objects, such as drifting wood, buoyant shells of organisms such as ''Spirula'', or man-made flotsam. Examples include goose barnacles and the bryozoan '' Jellyella''. By themselves these animals cannot float, which contrasts them with true planktonic organisms, such as '' Velella'' and the Portuguese Man o' War, which are buoyant. Pseudoplankton are often found in the guts of filtering zooplankters.

Tychoplankton

Tychoplankton are organisms, such as free-living or attached benthic organisms and other non-planktonic organisms, that are carried into the plankton through a disturbance of their benthic habitat, or by winds and currents. This can occur by direct turbulence or by disruption of the substrate and subsequent entrainment in the water column. Tychoplankton are, therefore, a primary subdivision for sorting planktonic organisms by duration of lifecycle spent in the plankton, as neither their entire lives nor particular reproductive portions are confined to planktonic existence. Tychoplankton are sometimes called ''accidental plankton''.

Mineralized plankton

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