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Zooplankton

Zooplankton sample including several species of copepods (1–5), gastropod larva (6) doliolids (7), fish eggs (8), and decapod larva (9) (Photo by Iole Di Capua)

Zooplankton are the heterotrophic component of the planktonic community (the "zoo-" prefix comes from Ancient Greek: ζῷον, romanizedzôion, lit. 'animal'), 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 Ancient Greek: φῠτόν, romanized: phutón, lit.'plant', 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 sunlight — in 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.[1]

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 (/ˈz.əplæŋktən/;[2] /ˌz.əˈplæŋktən/)[3] are heterotrophic (sometimes detritivorous) plankton. The word zooplankton is derived from Ancient Greek: ζῷον, romanizedzôion, lit. 'animal'; and πλᾰγκτός, planktós, 'wanderer; drifter'.[4]

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.[5] 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.[5] 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.[5]

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,[clarification needed] for instance, during the spring bloom. Zooplankton are also a key link in the biomagnification of pollutants such as mercury.[6]

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.[8]

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.[9][10] It has a paramount effect on growth, reproduction, feeding strategies and mortality.[11] 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.[12][13]

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.[14] Body size is sensitive to changes in temperature due to the thermal dependence of physiological processes.[15] 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.[16] 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.[17] This pattern of body size variation, known as the temperature-size rule (TSR),[18] has been observed for a wide range of ectotherms, including single-celled and multicellular species, invertebrates and vertebrates.[17][19][13]

The processes underlying the inverse relationship between body size and temperature remain to be identified.[17] 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.[20][21][13]

Zooplankton can be broken down into size classes[22] 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.[23] 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.[24][25] Microzooplankton are also pivotal regenerators of nutrients which fuel primary production and food sources for metazoans.[25][26]

Despite their ecological importance, microzooplankton remain understudied. Routine oceanographic observations seldom monitor microzooplankton biomass or herbivory rate, although