Zooplankton

Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequently, they drift or are carried along by currents in the ocean, or by currents in seas, lakes or rivers.

Zooplankton can be contrasted with phytoplankton (cyanobacteria and microalgae), which are the plant-like component of the plankton community (the " phyto-" prefix comes from , although taxonomically ''not'' plants). Zooplankton are heterotrophic (other-feeding), whereas phytoplankton are autotrophic (self-feeding), often generating biological energy and macromolecules through chlorophyllic carbon fixation using sunlightin other words, zooplankton cannot manufacture their own food, while phytoplankton can. As a result, zooplankton must acquire nutrients by feeding on other organisms such as phytoplankton, which are generally smaller than zooplankton. Most zooplankton are microscopic but some (such as jellyfish) are macroscopic, meaning they can be seen with the naked eye.

Many protozoans (single-celled protists that prey on other microscopic life) are zooplankton, including zooflagellates, foraminiferans, radiolarians, some dinoflagellates and marine microanimals. Macroscopic zooplankton include pelagic cnidarians, ctenophores, molluscs, arthropods and tunicates, as well as planktonic arrow worms and bristle worms.

The distinction between autotrophy and heterotrophy often breaks down in very small organisms. Recent studies of marine microplankton have indicated over half of microscopic plankton are mixotrophs, which can obtain energy and carbon from a mix of internal plastids and external sources. Many marine microzooplankton are mixotrophic, which means they could also be classified as phytoplankton.

Overview

Zooplankton (; ) are heterotrophic (sometimes detritivorous) plankton. The word ''zooplankton'' is derived from ; and .

Zooplankton is a categorization spanning a range of organism sizes including small protozoans and large metazoans. It includes holoplanktonic organisms whose complete life cycle lies within the plankton, as well as meroplanktonic organisms that spend part of their lives in the plankton before graduating to either the nekton or a sessile, benthic existence. Although zooplankton are primarily transported by ambient water currents, many have locomotion, used to avoid predators (as in diel vertical migration) or to increase prey encounter rate.

Just as any species can be limited within a geographical region, so are zooplankton. However, species of zooplankton are not dispersed uniformly or randomly within a region of the ocean. As with phytoplankton, 'patches' of zooplankton species exist throughout the ocean. Though few physical barriers exist above the mesopelagic, specific species of zooplankton are strictly restricted by salinity and temperature gradients, while other species can withstand wide temperature and salinity gradients. Zooplankton patchiness can also be influenced by biological factors, as well as other physical factors. Biological factors include breeding, predation, concentration of phytoplankton, and vertical migration. The physical factor that influences zooplankton distribution the most is mixing of the water column (upwelling and downwelling along the coast and in the open ocean) that affects nutrient availability and, in turn, phytoplankton production.

Through their consumption and processing of phytoplankton and other food sources, zooplankton play a role in aquatic food webs, as a resource for consumers on higher trophic levels (including fish), and as a conduit for packaging the organic material in the biological pump. Since they are typically small, zooplankton can respond rapidly to increases in phytoplankton abundance, for instance, during the spring bloom. Zooplankton are also a key link in the biomagnification of pollutants such as mercury.

File:Typical ocean models featuring zooplankton 2.jpg,

, alt=Upper left: Biogeochemical models                        Right: Ecosystem models      Lower left: Size-spectra modelsThese models also have temporal and spatial components.

Ecologically important protozoan zooplankton groups include the foraminiferans, radiolarians and dinoflagellates (the last of these are often mixotrophic). Important metazoan zooplankton include cnidarians such as jellyfish and the Portuguese Man o' War; crustaceans such as cladocerans, copepods, ostracods, isopods, amphipods, mysids and krill; chaetognaths (arrow worms); molluscs such as pteropods; and chordates such as salps and juvenile fish. This wide phylogenetic range includes a similarly wide range in feeding behavior: filter feeding, predation and symbiosis with autotrophic phytoplankton as seen in corals. Zooplankton feed on bacterioplankton, phytoplankton, other zooplankton (sometimes cannibalistically), detritus (or marine snow) and even nektonic organisms. As a result, zooplankton are primarily found in surface waters where food resources (phytoplankton or other zooplankton) are abundant.

Zooplankton can also act as a disease reservoir. Crustacean zooplankton have been found to house the bacterium ''Vibrio cholerae'', which causes cholera, by allowing the cholera vibrios to attach to their chitinous exoskeletons. This symbiotic relationship enhances the bacterium's ability to survive in an aquatic environment, as the exoskeleton provides the bacterium with carbon and nitrogen.

Size classification

Body size has been defined as a "master trait" for plankton as it is a morphological characteristic shared by organisms across taxonomy that characterises the functions performed by organisms in ecosystems. It has a paramount effect on growth, reproduction, feeding strategies and mortality. One of the oldest manifestations of the biogeography of traits was proposed over 170 years ago, namely Bergmann's rule, in which field observations showed that larger species tend to be found at higher, colder latitudes. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License

In the oceans, size is critical in determining trophic links in planktonic ecosystems and is thus a critical factor in regulating the efficiency of the biological carbon pump. Body size is sensitive to changes in temperature due to the thermal dependence of physiological processes. The plankton is mainly composed of ectotherms which are organisms that do not generate sufficient metabolic heat to elevate their body temperature, so their metabolic processes depends on external temperature. Consequently, ectotherms grow more slowly and reach maturity at a larger body size in colder environments, which has long puzzled biologists because classic theories of life-history evolution predict smaller adult sizes in environments delaying growth. This pattern of body size variation, known as the temperature-size rule (TSR), has been observed for a wide range of ectotherms, including single-celled and multicellular species, invertebrates and vertebrates.

The processes underlying the inverse relationship between body size and temperature remain to be identified. Despite temperature playing a major role in shaping latitudinal variations in organism size, these patterns may also rely on complex interactions between physical, chemical and biological factors. For instance, oxygen supply plays a central role in determining the magnitude of ectothermic temperature-size responses, but it is hard to disentangle the relative effects of oxygen and temperature from field data because these two variables are often strongly inter-related in the surface ocean.

Zooplankton can be broken down into size classes which are diverse in their morphology, diet, feeding strategies, etc. both within classes and between classes:

Microzooplankton

Microzooplankton are defined as heterotrophic and mixotrophic plankton. They primarily consist of phagotrophic protists, including ciliates, dinoflagellates, and mesozooplankton nauplii. Microzooplankton are major grazers of the plankton community. As the primary consumers of marine phytoplankton, microzooplankton consume ~ 59–75% daily of the marine primary production, much larger than mesozooplankton. That said, macrozooplankton can sometimes have greater consumption rates in eutrophic ecosystems because the larger phytoplankton can be dominant there. Microzooplankton are also pivotal regenerators of nutrients which fuel primary production and food sources for metazoans. Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.

Despite their ecological importance, microzooplankton remain understudied. Routine oceanographic observations seldom monitor microzooplankton biomass or herbivory rate, although the dilution technique, an elegant method of measuring microzooplankton herbivory rate, has been developed for over four decades (Landry and Hassett 1982). The number of observations of microzooplankton herbivory rate is around 1600 globally, far less than that of primary productivity (> 50,000). This makes validating and optimizing the grazing function of microzooplankton difficult in ocean ecosystem models.

Mesozooplankton

Mesozooplankton are one of the larger size classes of zooplankton. In most regions, mesozooplankton are dominated by copepods, such as Calanus finmarchicus and Calanus helgolandicus. Mesozooplankton are an important prey for fish.

As plankton are rarely fished, it has been argued that mesoplankton abundance and species composition can be used to study marine ecosystems' response to climate change. This is because they have life cycles that generally last less than a year, meaning they respond to climate changes between years. Sparse, monthly sampling will still indicate vacillations.

Taxonomic groups

Protozoans

Protozoans are protists that feed on organic matter such as other microorganisms or organic tissues and debris. Historically, the protozoa were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae. Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to identify single-celled organisms that can move independently and feed by heterotrophy.

Marine protozoans include zooflagellates, foraminiferans, radiolarians and some dinoflagellates.

Radiolarians

Radiolarians are unicellular predatory protists encased in elaborate globular shells usually made of silica and pierced with holes. Their name comes from the Latin for "radius". They catch prey by extending parts of their body through the holes. As with the silica frustules of diatoms, radiolarian shells can sink to the ocean floor when radiolarians die and become preserved as part of the ocean sediment. These remains, as microfossils, provide valuable information about past oceanic conditions.Wassilieff, Maggy (2006
"Plankton – Animal plankton"
''Te Ara – the Encyclopedia of New Zealand''. Accessed: 2 November 2019.

File:Mikrofoto.de-Radiolarien 6.jpg, Like diatoms, radiolarians come in many shapes
File:Podocyrtis papalis Ehrenberg - Radiolarian (30448963206).jpg, Also like diatoms, radiolarian shells are usually made of silicate
File:Acantharian radiolarian Xiphacantha (Haeckel).jpg, However acantharian radiolarians have shells made from strontium sulfate crystals
File:Spherical radiolarian 2.jpg, Cutaway schematic diagram of a spherical radiolarian shell

Foraminiferans

Like radiolarians, foraminiferans (''forams'' for short) are single-celled predatory protists, also protected with shells that have holes in them. Their name comes from the Latin for "hole bearers". Their shells, often called tests, are chambered (forams add more chambers as they grow). The shells are usually made of calcite, but are sometimes made of agglutinated sediment particles or chiton, and (rarely) silica. Most forams are benthic, but about 40 species are planktic. They are widely researched with well-established fossil records which allow scientists to infer a lot about past environments and climates.

File:EB1911 Foraminifera - Section of Rotalia beccarii.jpg, section showing chambers of a spiral foram
File:Live Ammonia tepida.jpg, Live '' Ammonia tepida'' streaming granular ectoplasm for catching food
File:Planktic Foraminifera of the northern Gulf of Mexico.jpg, Group of planktonic forams
File:All Gizah Pyramids.jpg, The Egyptian pyramids were constructed from limestone that contained nummulites.

Amoeba

File:Amoeba proteus 2.jpg, Naked amoeba sketch showing food vacuoles and ingested diatom
File:Arcella sp.jpg, Shell or test of a testate amoeba, '' Arcella'' sp.
File:Collection Penard MHNG Specimen 533-2-1 Pamphagus granulatus.tif, Xenogenic testate amoeba covered in diatoms

Ciliates

File:Stylonychia putrina - 160x - II (13215594964).jpg, '' Stylonychia putrina''
File:Holophyra ovum - 400x (9836710085).jpg, ''Holophyra'' ovum
File:Mikrofoto.de-Blepharisma japonicum 15.jpg, '' Blepharisma japonicum''
File:The ciliate Frontonia sp.jpg, This ciliate is digesting cyanobacteria. The mouth is at the bottom right.

Dinoflagellates

Dinoflagellates are a phylum of unicellular flagellates with about 2,000 marine species. Some dinoflagellates are predatory, and thus belong to the zooplankton community. Their name comes from the Greek "dinos" meaning ''whirling'' and the Latin "flagellum" meaning a ''whip'' or ''lash''. This refers to the two whip-like attachments (flagella) used for forward movement. Most dinoflagellates are protected with red-brown, cellulose armour. Excavates may be the most basal flagellate lineage.

File:Gyrodinium dinoflagellate.jpg, '' Gyrodinium'', one of the few naked dinoflagellates which lack armour
File:Protoperidinium dinoflagellate.jpg, The dinoflagellate ''Protoperidinium'' extrudes a large feeding veil to capture prey
File:Radiolarian - Podocyrtis (Lampterium) mitra Ehrenberg - 160x.jpg, Nassellarian radiolarians can be in symbiosis with dinoflagellates

Dinoflagellates often live in symbiosis with other organisms. Many nassellarian radiolarians house dinoflagellate symbionts within their tests. The nassellarian provides ammonium and carbon dioxide for the dinoflagellate, while the dinoflagellate provides the nassellarian with a mucous membrane useful for hunting and protection against harmful invaders. There is evidence from DNA analysis that dinoflagellate symbiosis with radiolarians evolved independently from other dinoflagellate symbioses, such as with foraminifera.

File:Ceratium tripos.jpg, ''Tripos muelleri'' is recognisable by its U-shaped horns
File:Archives de zoologie expérimentale et générale (1920) (20299351186).jpg, '' Oodinium'', a genus of parasitic dinoflagellates, causes velvet disease in fish
File:Karenia brevis.jpg, '' Karenia brevis'' produces red tides highly toxic to humans
File:Algal bloom(akasio) by Noctiluca in Nagasaki.jpg, Red tide

Mixotrophs

A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comprise more than half of all microscopic plankton. There are two types of eukaryotic mixotrophs: those with their own chloroplasts, and those with endosymbionts—and others that acquire them through kleptoplasty or by enslaving the entire phototrophic cell.

The distinction between plants and animals often breaks down in very small organisms. Possible combinations are photo- and chemotrophy, litho- and organotrophy, auto- and heterotrophy or other combinations of these. Mixotrophs can be either eukaryotic or prokaryotic. They can take advantage of different environmental conditions.

Many marine microzooplankton are mixotrophic, which means they could also be classified as phytoplankton. Recent studies of marine microzooplankton found 30–45% of the ciliate abundance was mixotrophic, and up to 65% of the amoeboid, foram and radiolarian biomass was mixotrophic.

''Phaeocystis'' species are endosymbionts to acantharian radiolarians. '' Phaeocystis'' is an important algal genus found as part of the marine phytoplankton around the world. It has a polymorphic life cycle, ranging from free-living cells to large colonies. It has the ability to form floating colonies, where hundreds of cells are embedded in a gel matrix, which can increase massively in size during blooms. As a result, ''Phaeocystis'' is an important contributor to the marine carbon and sulfur cycles.

File:Tintinnid ciliate Favella.jpg, Tintinnid ciliate ''Favella''
File:Euglena mutabilis - 400x - 1 (10388739803) (cropped).jpg, '' Euglena mutabilis'', a photosynthetic flagellate
File:Stichotricha secunda - 400x (14974779356).jpg, Zoochlorellae (green) living inside the ciliate ''Stichotricha secunda''
File:Dinophysis acuta.jpg, The dinoflagellate ''Dinophysis acuta''

A number of forams are mixotrophic. These have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates. Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters. Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.

By trophic orientation, dinoflagellates are all over the place. Some dinoflagellates are known to be photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey ( phagotrophy). Some species are endosymbionts of marine animals and other protists, and play an important part in the biology of coral reefs. Others predate other protozoa, and a few forms are parasitic. Many dinoflagellates are mixotrophic and could also be classified as phytoplankton. The toxic dinoflagellate '' Dinophysis acuta'' acquire chloroplasts from its prey. "It cannot catch the cryptophytes by itself, and instead relies on ingesting ciliates such as the red '' Myrionecta rubra'', which sequester their chloroplasts from a specific cryptophyte clade (Geminigera/Plagioselmis/Teleaulax)".

Metazoa (animals)

Free-living species in the crustacean class Copepoda are typically 1 to 2 mm long with teardrop-shaped bodies. Like all crustaceans, their bodies are divided into three sections: head, thorax, and abdomen, with two pairs of antennae; the first pair is often long and prominent. They have a tough exoskeleton made of calcium carbonate and usually have a single red eye in the centre of their transparent head. About 13,000 species of copepods are known, of which about 10,200 are marine. They are usually among the more dominant members of the zooplankton.

In addition to copepods the crustacean classes ostracods, branchiopods and malacostracans also have planktonic members. Barnacles are planktonic only during the larval stage.

Metazoan