Zooplankton

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Sample of zooplankton which includes fish eggs, doliolids, several species of copepods, gastropod

and decapod larva

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Plankton

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Trophic groups

 Phytoplankton
 Zooplankton
 Mixoplankton
 Mycoplankton
 Bacterioplankton
  Virioplankton

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Size groups

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Taxonomic groups

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Other groups

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Zooplankton (/ˈzoʊ.əˌplæŋktən, ˈzuː(ə)-, ˈzoʊoʊ-/,[1] /ˌzoʊ.əˈplæŋktən, -tɒn/)[2] are 
heterotrophic (sometimes detritivorous) plankton (cf. phytoplankton). Plankton
are organisms drifting in oceans, seas, and bodies of fresh water. The
word zooplankton is derived from the Greek zoon (ζῴον), meaning "animal",
and planktos (πλαγκτός), meaning "wanderer" or "drifter". [3] Individual zooplankton
are usually microscopic, but some (such as jellyfish) are larger and visible to
the naked eye.

Contents

 1Overview
 2Classification by size
 3Taxonomic groups
o 3.1Protozoa
 3.1.1Radiolarians
 3.1.2Foraminiferans
 3.1.3Amoeba
 3.1.4Ciliates
 3.1.5Dinoflagellates
o 3.2Mixotrophs
o 3.3Metazoa
 3.3.1Holoplankton and meroplankton
 3.3.2Ichthyoplankton
 3.3.3Gelatinous zooplankton
 4Microzooplankton
 5Role in food webs
 6Role in biogeochemistry
o 6.1Sloppy feeding and release of DOM
o 6.2Carbon Export
 7See also
 8References
 9External links

Overview[edit]
Zooplankton are the animal component of the planktonic community ("zoo"
comes from the Greek word for animal). They are heterotrophic (other-feeding),
meaning they cannot produce their own food and must consume instead other
plants or animals as food. In particular, this means they eat phytoplankton.
Zooplankton are generally larger than phytoplankton, mostly still microscopic but
some 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.
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.

 Typical models featuring zooplankton

 

      Upper left: Biogeochemical models                        Right: Ecosystem models

      Lower left: Size-spectra models

These models also have temporal and spatial components.[4]

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 copepods, ostracods, isopods, amphipods, mysids and krill; chaetognaths (arr
ow 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.
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]
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.[7]

Classification by size[edit]
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by adding to it. (March 2021)

Taxonomic groups[edit]
Protozoa[edit]
Further information: marine protists § Protozoans
Protozoans are protists that feed on organic matter such as
other microorganisms or organic tissues and debris.[8][9] 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.[10][11] 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[edit]
Radiolarian shapes

          Drawings by Haeckel 1904 (click for details)

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.[12]

Like diatoms, radiolarians come in many shapes

 

Also like diatoms, radiolarian shells are usually made of silicate

 

However acantharian radiolarians have shells made from strontium sulfate crystals

 
 

Cutaway schematic diagram of a spherical radiolarian shell

External video

 Radiolarian geometry

 Ernst Haeckel's radiolarian engravings

Foraminiferans[edit]
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) of silica. Most forams are benthic, but about 40 species are planktic.
[13]
 They are widely researched with well established fossil records which allow
scientists to infer a lot about past environments and climates. [12]
Foraminiferans

...can have more than one nucleus

 ...and defensive spines
Foraminiferans are important unicellular zooplankton protists, with calcium tests

section showing chambers of a spiral foram

 

Live Ammonia tepida streaming granular ectoplasm for catching food

 

Group of planktonic forams

  

The Egyptian pyramids were constructed from limestone that contained nummulites.[14]

External video

 foraminiferans

 Foraminiferal networks and growth

Amoeba[edit]
Shelled and naked amoeba

Testate amoeba, Cyphoderia sp.

Naked amoeba, Chaos sp.
                  Amoeba can be shelled (testate) or naked
 

Naked amoeba sketch showing food vacuoles and ingested diatom

 

Shell or test of a testate amoeba, Arcella sp.

 

Xenogenic testate amoeba covered in diatoms

Ciliates[edit]
 

Stylonychia putrina
 

Holophyra ovum
 

Blepharisma japonicum
 
 

This ciliate is digesting cyanobacteria. The mouth is at the bottom right.

Dinoflagellates[edit]
See also: Predatory dinoflagellate
Dinoflagellates are part of the algae group, and form a phylum of unicellular
flagellates with about 2,000 marine species. [15] The 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. Like other
phytoplankton, dinoflagellates are r-strategists which under right conditions
can bloom and create red tides. Excavates may be the most basal flagellate
lineage.[16]
Dinoflagellates

        Armoured

        Unarmoured
Traditionally dinoflagellates have been presented as armoured or unarmoured
 

Gyrodinium, one of the few naked dinoflagellates which lack armour

 

The dinoflagellate Protoperidinium extrudes a large feeding veil to capture prey

 

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.
[17]
 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. [18] There is evidence
from DNA analysis that dinoflagellate symbiosis with radiolarians evolved
independently from other dinoflagellate symbioses, such as with foraminifera.[19]
 

Tripos muelleri is recognisable by its U-shaped horns

 

Oodinium, a genus of parasitic dinoflagellates, causes velvet disease in fish[20]

 

Karenia brevis produces red tides highly toxic to humans[21]

 

Red tide
Mixotrophs[edit]
See also: Mixotroph and Mixotrophic dinoflagellate
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. [22] 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. [23]
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.
[24]
 They can take advantage of different environmental conditions. [25]
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.[26]

Mixotrophic zooplankton that combine phototrophy and heterotrophy – table based on Stoecker et. al.,
2017 [27]

Description Example Further examples

Called nonconstitutive mixotrophs by Mitra et. al., 2016.[28] Zooplankton that are photosynthetic:

microzooplankton or metazoan zooplankton that acquire phototrophy through chloroplast retention a or
maintenance of algal endosymbionts.

Protists that
retain
chloroplasts and
Generalists Most oligotrich ciliates that retain plastidsa
rarely other
organelles from
many algal taxa
 1. Protists that
retain
chloroplasts and
sometimes other
Dinophysis Dinophysis spp.
organelles from
acuminata Myrionecta rubra
one algal species
or very closely
related algal
species
Specialists
2. Protists or
zooplankton
with algal Metazooplankton with algal endosymbionts
endosymbionts Most
Noctiluca
of only one mixotrophic Rhizaria (Acantharea, Polycystinea,
scintillans
algal species and Foraminifera)
or very closely Green Noctiluca scintillans
related algal
species

Chloroplast (or plastid) retention = sequestration = enslavement. Some plastid-retaining species also retain other organelles and prey
a

cytoplasm.

Phaeocystis species are endosymbionts to acantharian radiolarians.[29]

[30]
 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.[31] 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.[32] As a result, Phaeocystis is an important
contributor to the marine carbon[33] and sulfur cycles.[34]

 Mixoplankton
 

Tintinnid ciliate Favella
 

Euglena mutabilis, a photosynthetic flagellate

 

Zoochlorellae (green) living inside the ciliate Stichotricha secunda

 
 

The dinoflagellate Dinophysis acuta

Mixotrophic radiolarians

Acantharian radiolarian hosts Phaeocystis symbionts

White Phaeocystis algal foam washing up on a beach

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.[13] Mixotrophic
foraminifers are particularly common in nutrient-poor oceanic waters. [35] Some
forams are kleptoplastic, retaining chloroplasts from ingested algae to
conduct photosynthesis.[36]
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).
[37]
 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)". [27]
Metazoa[edit]

Octopus larva and pteropod

Copepods are typically 1 to 2 mm long with a 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. [38] About 13,000 species of
copepods are known, of which about 10,200 are marine. [39][40] They are usually
among the more dominant members of the zooplankton. [41]

 Metazoan zooplankton


Copepod with eggs
 

Jellyfish
  

Segmented worm
 

Amphipod
 

Krill
 
 

Blue ocean slug

Holoplankton and meroplankton[edit]
Ichthyoplankton[edit]

 Ichthyoplankton


Salmon eggs
 

Juvenile planktonic squid
 
 

Ocean sunfish larvae (2.7mm)

 

Boxfish larva
Gelatinous zooplankton[edit]
Gelatinous zooplankton include ctenophores, medusae, 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 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.[42][43] That view has recently been
challenged. Jellyfish, and more gelatinous zooplankton in general, which
include salps and ctenophores, 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. [42] But jellyfish bloom in vast numbers, and it
has been shown they form major components in the diets
of tuna, spearfish and swordfish as well as various birds and invertebrates such
as octopus, sea cucumbers, crabs and amphipods.[44][43] "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."[43] According to a 2017 study, narcomedusae consume the greatest
diversity of mesopelagic prey, followed
by physonect siphonophores, ctenophores and cephalopods.[45] 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.[45]

Microzooplankton[edit]
Microzooplankton: major grazers of the plankton...

Role in food webs[edit]

 Pelagic food web


Pelagic food web and the biological pump. Links among the ocean's biological pump and
pelagic food web and the ability to sample these components remotely from ships, satellites,
and autonomous vehicles. Light blue waters are the euphotic zone, while the darker blue
waters represent the twilight zone.[46]

Role in biogeochemistry[edit]
In addition to linking primary producers to higher trophic levels 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 fecal
pellets, 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.[47]
Sloppy feeding and release of DOM[edit]
Excretion and sloppy feeding (the physical breakdown of food source) make up
80% and 20% of crustacean zooplankton-mediated DOM release respectively.
[48]
 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.[49][50] 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.
[47]
 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.[51][52] 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.[49]
Comparison of zooplankton-mediated carbon cycles [53]

Kerguelen Plateau
Naturally iron-fertilized
On the Kerguelen Plateau in summer, high iron levels lead to high chlorophyll a as a proxy for algae
biomass at the surface. The diverse zooplankton community feeds on the sinking particle flux and acts
as a gate-keeper to the deeper ocean by ingesting and fragmenting sinking particles and,
consequently, significantly reducing the export flux out of the epipelagic. The main export particles
are diatom resting spores, which bypass the intense grazing pressure, followed by fecal pellets. [53]

Southern Ocean waters

High nutrient, low chlorophyll
In Southern Ocean waters in summer, iron levels are relatively low and support a more diverse
phytoplankton community, but with lower biomass, which, in turn, affects zooplankton community
composition and biomass. The grazing pressure during summer is focused mostly on picoplankton,
which leaves large particles for export. [53]
Grazing and fragmentation of particles at both sites increases nutrient recycling in the upper water column
 Sloppy feeding by zooplankton
DOC = dissolved organic carbon
POC = particulate organic carbon
Adapted from Møller et al. (2005),[54]

Saba et al. (2009)[55] and Steinberg et al. (2017). [47]

Carbon Export[edit]
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 microbial
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. [56] Carcasses are also gaining recognition as
being important contributors to carbon export. Jelly falls – 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 benthic organisms. [