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Zooplankton are the component of the community (the "" 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 , or .
Zooplankton can be contrasted with phytoplankton (cyanobacteria and microalgae), which are the plant-like component of the plankton community (the "" prefix comes from , although taxonomically not ). Zooplankton are heterotrophic (other-feeding), whereas phytoplankton are autotrophic (self-feeding), often generating biological energy and through carbon fixation using other words, zooplankton cannot manufacture their own food, while phytoplankton can. As a result, zooplankton must acquire by feeding on other 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 Marine protists that prey on other microscopic life) are zooplankton, including , , , some and marine microanimals. Macroscopic zooplankton include pelagic , ctenophores, , and , as well as planktonic chaetognatha and Polychaete.
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 , 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.
Zooplankton is a categorization spanning a range of organism sizes including small and large . It includes organisms whose complete life cycle lies within the plankton, as well as organisms that spend part of their lives in the plankton before graduating to either the nekton or a sessile, benthos 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. (1993). 9780750633840, Elsevier. ISBN 9780750633840 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 , as a resource for consumers on higher (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 such as mercury.
|alt=Upper left: Biogeochemical models Right: Ecosystem models Lower left: Size-spectra modelsThese models also have temporal and spatial components.
Ecology important protozoan zooplankton groups include the , and (the last of these are often ). Important metazoan zooplankton include such as jellyfish and the Portuguese Man o' War; such as , , , , , mysids and krill; Chaetognatha (arrow worms); Mollusca such as ; and such as and juvenile fish. This wide phylogeny range includes a similarly wide range in feeding behavior: filter feeder, predation and symbiosis with autotrophic phytoplankton as seen in corals. Zooplankton feed on bacterioplankton, phytoplankton, other zooplankton (sometimes Cannibalism), detritus (or marine snow) and even Nekton. 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 . This symbiotic relationship enhances the bacterium's ability to survive in an aquatic environment, as the exoskeleton provides the bacterium with carbon and nitrogen.
In the oceans, size is critical in determining Trophic level 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 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), (1994). 9780120139255 ISBN 9780120139255 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:
| picozooplankton | 2μm |
| nanozooplankton | 2–20μm |
| microzooplankton | 20–200μm |
| mesozooplankton | 0.2–20 millimeters |
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.
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.
Marine protozoans include , , and some .
Dinoflagellates often live in symbiosis with other organisms. Many nassellarian radiolarians house dinoflagellate Symbiosis within their tests. (2025). 9783319281476, Springer, Cham. ISBN 9783319281476 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.
The distinction between plants and animals often breaks down in very small organisms. Possible combinations are phototroph and , lithotroph and , autotroph and or other combinations of these. Mixotrophs can be either eukaryote or prokaryote. 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.
| Called nonconstitutive mixotrophs by Mitra et al., 2016. Zooplankton that are photosynthetic: microzooplankton or metazoan zooplankton that acquire phototrophy through chloroplast retentiona or maintenance of algal endosymbionts. | ||||||
| Generalists | Protists that retain chloroplasts and rarely other organelles from many algal taxa | Most oligotrich ciliates that retain plastidsa | ||||
| Specialists | 1. Protists that retain chloroplasts and sometimes other organelles from one algal species or very closely related algal species | Dinophysis acuminata | Dinophysis spp. Myrionecta rubra | |||
| 2. Protists or zooplankton with algal endosymbionts of only one algal species or very closely related algal species | Noctiluca scintillans | with algal Most mixotrophic Rhizaria (Acantharea, Polycystinea, and Foraminifera) Green Noctiluca scintillans | ||||
| aChloroplast (or plastid) retention = sequestration = enslavement. Some plastid-retaining species also retain other organelles and prey cytoplasm. | ||||||
Phaeocystis species are endosymbionts to Acantharea 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 Algal bloom. As a result, Phaeocystis is an important contributor to the marine Carbon cycle and .
A number of forams are mixotrophic. These have unicellular algae as , from diverse lineages such as the green algae, red algae, golden algae, , and . Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters. Some forams are kleptoplasty, retaining from ingested algae to conduct photosynthesis.
By trophic orientation, dinoflagellates are all over the place. Some dinoflagellates are known to be photosynthesis, but a large fraction of these are in fact mixotrophy, combining photosynthesis with ingestion of prey (phagotrophy). Some species are of marine animals and other protists, and play an important part in the biology of . 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)".
In addition to copepods the crustacean classes , Branchiopoda and also have planktonic members. are planktonic only during the larval stage. Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea
That view has recently been challenged. Jellyfish, and more gelatinous zooplankton in general, which include and , are very diverse, fragile with no hard parts, difficult to see and monitor, subject to rapid population swings and often live inconveniently far from shore or deep in the ocean. It is difficult for scientists to detect and analyse jellyfish in the guts of predators, since they turn to mush when eaten and are rapidly digested. But jellyfish bloom in vast numbers, and it has been shown they form major components in the diets of tuna, Tetrapturus and swordfish as well as various birds and invertebrates such as octopus, , and .Cardona, L., De Quevedo, I.Á., Borrell, A. and Aguilar, A. (2012) "Massive consumption of gelatinous plankton by Mediterranean apex predators". PLOS ONE, 7(3): e31329. "Despite their low energy density, the contribution of jellyfish to the energy budgets of predators may be much greater than assumed because of rapid digestion, low capture costs, availability, and selective feeding on the more energy-rich components. Feeding on jellyfish may make marine predators susceptible to ingestion of plastics." According to a 2017 study, narcomedusae consume the greatest diversity of mesopelagic prey, followed by physonect , and .
The importance of the so-called "jelly web" is only beginning to be understood, but it seems medusae, ctenophores and siphonophores can be key predators in deep pelagic food webs with ecological impacts similar to predator fish and squid. Traditionally gelatinous predators were thought ineffectual providers of marine trophic pathways, but they appear to have substantial and integral roles in deep pelagic food webs.Choy, C.A., Haddock, S.H. and Robison, B.H. (2017) "Deep pelagic food web structure as revealed by in situ feeding observations". Proceedings of the Royal Society B: Biological Sciences, 284(1868): 20172116. . Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
Grazing is a central, rate-setting process in ocean ecosystems and a driver of marine biogeochemical cycling. In all ocean ecosystems, grazing by heterotrophic protists constitutes the single largest loss factor of marine primary production and alters particle size distributions. Grazing affects all pathways of export production, rendering grazing important both for surface and deep carbon processes. Predicting central paradigms of ocean ecosystem function, including responses to environmental change requires accurate representation of grazing in global biogeochemical, ecosystem and cross-biome-comparison models. Several large-scale analyses have concluded that phytoplankton losses, which are dominated by grazing are the putative explanation for annual cycles in phytoplankton biomass, accumulation rates and export production.
Metazoa (animals)
Free-living species in the crustacean class 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.
Holoplankton and meroplankton
Ichthyoplankton
Ichthyoplankton are the Fish eggs and larvae of fish ("ichthyo" comes from the Greek word for fish). They are planktonic because 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 juvenile fish. 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.
Gelatinous zooplankton
Gelatinous zooplankton include , Jellyfish, salps, and Chaetognatha in coastal waters. Jellyfish are slow swimmers, and most species form part of the plankton. Traditionally jellyfish have been viewed as Trophic level dead ends, minor players in the marine food web, gelatinous organisms with a body plan largely based on water that offers little nutritional value or interest for other organisms apart from a few specialised predators such as the ocean sunfish and the leatherback sea turtle.Hamilton, G. (2016) "The secret lives of jellyfish: long regarded as minor players in ocean ecology, jellyfish are actually important parts of the marine food web". Nature, 531(7595): 432–435. Graeme Hays, Doyle, T.K. and Houghton, J.D. (2018) "A paradigm shift in the trophic importance of jellyfish?" Trends in ecology & evolution, 33(11): 874–884.
Role in food webs
Grazing by single-celled zooplankton accounts for the majority of organic carbon loss from marine primary production. Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License. However, zooplankton grazing remains one of the key unknowns in global predictive models of carbon flux, the marine food web structure and ecosystem characteristics, because empirical grazing measurements are sparse, resulting in poor parameterisation of grazing functions. To overcome this critical knowledge gap, it has been suggested that a focused effort be placed on the development of instrumentation that can link changes in phytoplankton biomass or optical properties with grazing.
Role in biogeochemistry
In addition to linking primary producers to higher in marine food webs, zooplankton also play an important role as "recyclers" of carbon and other nutrients that significantly impact marine biogeochemical cycles, including the biological pump. This is particularly important in the oligotrophic waters of the open ocean. Through sloppy feeding, excretion, egestion, and leaching of , zooplankton release dissolved organic matter (DOM) which controls DOM cycling and supports the microbial loop. Absorption efficiency, respiration, and prey size all further complicate how zooplankton are able to transform and deliver carbon to the deep ocean.
Sloppy feeding and release of DOM
Excretion and sloppy feeding (the physical breakdown of food source) make up 80% and 20% of crustacean zooplankton-mediated DOM release respectively. In the same study, fecal pellet leaching was found to be an insignificant contributor. For protozoan grazers, DOM is released primarily through excretion and egestion and gelatinous zooplankton can also release DOM through the production of mucus. Leaching of fecal pellets can extend from hours to days after initial egestion and its effects can vary depending on food concentration and quality. Various factors can affect how much DOM is released from zooplankton individuals or populations. Absorption efficiency (AE) is the proportion of food absorbed by plankton that determines how available the consumed organic materials are in meeting the required physiological demands. Depending on the feeding rate and prey composition, variations in AE may lead to variations in fecal pellet production, and thus regulates how much organic material is recycled back to the marine environment. Low feeding rates typically lead to high AE and small, dense pellets, while high feeding rates typically lead to low AE and larger pellets with more organic content. Another contributing factor to DOM release is respiration rate. Physical factors such as oxygen availability, pH, and light conditions may affect overall oxygen consumption and how much carbon is loss from zooplankton in the form of respired CO2. The relative sizes of zooplankton and prey also mediate how much carbon is released via sloppy feeding. Smaller prey are ingested whole, whereas larger prey may be fed on more "sloppily", that is more biomatter is released through inefficient consumption. There is also evidence that diet composition can impact nutrient release, with carnivorous diets releasing more dissolved organic carbon (DOC) and ammonium than omnivorous diets.
Carbon export
Zooplankton play a critical role in supporting the ocean's biological pump through various forms of carbon export, including the production of fecal pellets, mucous feeding webs, molts, and carcasses. Fecal pellets are estimated to be a large contributor to this export, with copepod size rather than abundance expected to determine how much carbon actually reaches the ocean floor. The importance of fecal pellets can vary both by time and location. For example, zooplankton bloom events can produce larger quantities of fecal pellets, resulting in greater measures of carbon export. Additionally, as fecal pellets sink, they are reworked by microbes in the water column, which can thus alter the carbon composition of the pellet. This affects how much carbon is recycled in the euphotic zone and how much reaches depth. Fecal pellet contribution to carbon export is likely underestimated; however, new advances in quantifying this production are currently being developed, including the use of isotopic signatures of amino acids to characterize how much carbon is being exported via zooplankton fecal pellet production. Carcasses are also gaining recognition as being important contributors to carbon export. – the mass sinking of gelatinous zooplankton carcasses – occur across the world as a result of large blooms. Because of their large size, these gelatinous zooplankton are expected to hold a larger carbon content, making their sinking carcasses a potentially important source of food for Benthos.
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