Plankton

Last updated

Marine microplankton and mesoplankton
Part of the contents of one dip of a hand net. The image contains diverse planktonic organisms, ranging from photosynthetic cyanobacteria and diatoms to many different types of zooplankton, including both holoplankton (permanent residents of the plankton) and meroplankton (temporary residents of the plankton, e.g., fish eggs, crab larvae, worm larvae). Marine microplankton.jpg
Part of the contents of one dip of a hand net. The image contains diverse planktonic organisms, ranging from photosynthetic cyanobacteria and diatoms to many different types of zooplankton, including both holoplankton (permanent residents of the plankton) and meroplankton (temporary residents of the plankton, e.g., fish eggs, crab larvae, worm larvae).

Plankton are the diverse collection of organisms that drift in water (or air) but are unable to actively propel themselves against currents (or wind). [1] [2] The individual organisms constituting plankton are called plankters. [3] In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish, and baleen whales.

Contents

Marine plankton include bacteria, archaea, algae, protozoa, microscopic fungi, [4] 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 lakes and rivers. Mostly, plankton just drift where currents take them, though some, like jellyfish, swim slowly but not fast enough to generally overcome the influence of currents.

Although plankton are usually thought of as inhabiting water, there are also airborne versions that live part of their lives drifting in the atmosphere. These aeroplankton include plant spores, pollen and wind-scattered seeds. They may also include 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. [5] This is because plankton are defined by their ecological niche and level of motility rather than by any phylogenetic or taxonomic classification. The "plankton" category differentiates these organisms from those that float on the water's surface, called neuston, those that can swim against a current, called nekton , and those that live on the deep sea floor, called benthos .

Terminology

Plankton (organisms that drift with water currents) can be contrasted with nekton (organisms that swim against water currents), neuston (organisms that live at the ocean surface) and benthos (organisms that live at the ocean floor). Neuston, Plankton, Nekton, Benthos.jpg
Plankton (organisms that drift with water currents) can be contrasted with nekton (organisms that swim against water currents), neuston (organisms that live at the ocean surface) and benthos (organisms that live at the ocean floor).

The name plankton was coined by German marine biologist Victor Hensen in 1887 from shortening the word halyplankton from Greek ᾰ̔́λςháls "sea" and πλανάωplanáō to "drift" or "wander". [6] :1 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. [7]

The amount and distribution of plankton depends on available nutrients, the state of water and a large amount of other plankton. [8]

The study of plankton is termed planktology and a planktonic individual is referred to as a plankter. [9] 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. [10]

Trophic groups

Plankton are primarily divided into broad functional (or trophic level) groups:

Mixoplankton

Recognition of the importance of mixotrophy as an ecological strategy is increasing, [15] as well as the wider role this may play in marine biogeochemistry. [16] Studies have shown that mixotrophs are much more important for marine ecology than previously assumed and comprise more than half of all microscopic plankton. [17] [18] Their presence acts as a buffer that prevents the collapse of ecosystems during times with little to no light. [19]

Size groups

Plankton species diversity
Diverse assemblages consist of unicellular and multicellular organisms with different sizes, shapes, feeding strategies, ecological functions, life cycle characteristics, and environmental sensitivities.
Courtesy of Christian Sardet/CNRS/Tara expeditions Plankton species diversity.jpg
Plankton species diversity
Diverse assemblages consist of unicellular and multicellular organisms with different sizes, shapes, feeding strategies, ecological functions, life cycle characteristics, and environmental sensitivities.
Courtesy of Christian Sardet/CNRS/Tara expeditions

Plankton are also often described in terms of size. Usually the following divisions are used: [21]

GroupSize range
    (ESD)
Examples
Megaplankton> 20 cm metazoans; e.g. jellyfish; ctenophores; salps and pyrosomes (pelagic Tunicata); Cephalopoda; Amphipoda
Macroplankton2→20 cm metazoans; e.g. Pteropoda; Chaetognaths; Euphausiacea (krill); Medusae; ctenophores; salps, doliolids and pyrosomes (pelagic Tunicata); Cephalopoda; Janthina and Recluzia (two genera of gastropods); Amphipoda
Mesoplankton0.2→20 mm metazoans; e.g. copepods; Medusae; Cladocera; Ostracoda; Chaetognaths; Pteropoda; Tunicata
Microplankton20→200 μm large eukaryotic protists; most phytoplankton; Protozoa Foraminifera; tintinnids; other ciliates; Rotifera; juvenile metazoansCrustacea (copepod nauplii)
Nanoplankton2→20 μmsmall eukaryotic protists; small diatoms; small flagellates; Pyrrophyta; Chrysophyta; Chlorophyta; Xanthophyta
Picoplankton 0.2→2 μmsmall eukaryotic protists; bacteria; Chrysophyta
Femtoplankton< 0.2 μm marine viruses

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. [22]

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

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. [24] 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. [25] [26] 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. [27] When airborne, these microbes can be transported long distances to coastal regions. If they hit land they can have an effect on animal, vegetation and human health. [28] 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%). [29] [30] [31] These aerosols are able to remain suspended in the atmosphere for about 31 days. [32] 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. [33] 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). [33]

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 [34]

Other groups

Gelatinous zooplankton

Jellyfish are gelatinous zooplankton. Jellyfish swarm.jpg
Jellyfish are 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. [36] Gelatinous zooplankton are often transparent. [37] 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. [38]

Ichthyoplankton

Salmon egg hatching into a sac fry. In a few days, the sac fry will absorb the yolk sac and start feeding on smaller plankton. Salmonlarvakils 2.jpg
Salmon egg hatching into a sac fry. In a few days, the sac fry will absorb the yolk sac and start feeding on smaller plankton.

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 food supply. Both eggs and larvae are themselves eaten by larger animals. [39] [40] Fish can produce high numbers of eggs which are often released into the open water column. Fish eggs typically have a diameter of about 1 millimetre (0.039 in). 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

Tomopteris, a holoplanktic bioluminescence polychaete worm Tomopteriskils.jpg
Tomopteris , a holoplanktic bioluminescence polychaete worm

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. [42] Holoplankton include both phytoplankton and zooplankton and vary in size. The most common plankton are protists. [43]

Meroplankton

Larva stage of a spiny lobster Larva de phyllosoma.jpg
Larva stage of a spiny lobster

Meroplankton are a wide variety of aquatic organisms that have both planktonic and benthic stages in their life cycles. Much of the meroplankton consists of larval stages of larger organisms. [34] Meroplankton can be contrasted with holoplankton, which are planktonic organisms that stay in the pelagic zone as plankton throughout their entire life cycle. [44] After some 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. [45] The planktonic larval stage is particularly crucial to many benthic invertebrates 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 hours to months. [34]

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. [46]

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. [47] This can occur by direct turbulence or by disruption of the substrate and subsequent entrainment in the water column. [47] [48] 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. [49] Tychoplankton are sometimes called accidental plankton.

Mineralized plankton

Distribution

World concentrations of surface ocean chlorophyll as viewed by satellite during the northern spring, averaged from 1998 to 2004. Chlorophyll is a marker for the distribution and abundance of phytoplankton. Plankton satellite image.jpg
World concentrations of surface ocean chlorophyll as viewed by satellite during the northern spring, averaged from 1998 to 2004. Chlorophyll is a marker for the distribution and abundance of phytoplankton.

Apart from aeroplankton, plankton inhabits oceans, seas, lakes and ponds. Local abundance varies horizontally, vertically and seasonally. The primary cause of this variability is the availability of light. All plankton ecosystems are driven by the input of solar energy (but see chemosynthesis), confining primary production to surface waters, and to geographical regions and seasons having abundant light.

A secondary variable is nutrient availability. Although large areas of the tropical and sub-tropical oceans have abundant light, they experience relatively low primary production because they offer limited nutrients such as nitrate, phosphate and silicate. This results from large-scale ocean circulation and water column stratification. In such regions, primary production usually occurs at greater depth, although at a reduced level (because of reduced light).

Despite significant macronutrient concentrations, some ocean regions are unproductive (so-called HNLC regions). [50] The micronutrient iron is deficient in these regions, and adding it can lead to the formation of phytoplankton algal blooms. [51] Iron primarily reaches the ocean through the deposition of dust on the sea surface. Paradoxically, oceanic areas adjacent to unproductive, arid land thus typically have abundant phytoplankton (e.g., the eastern Atlantic Ocean, where trade winds bring dust from the Sahara Desert in north Africa).

While plankton are most abundant in surface waters, they live throughout the water column. At depths where no primary production occurs, zooplankton and bacterioplankton instead consume organic material sinking from more productive surface waters above. This flux of sinking material, so-called marine snow, can be especially high following the termination of spring blooms.

The local distribution of plankton can be affected by wind-driven Langmuir circulation and the biological effects of this physical process.

Ecological significance

Food chain

External videos
Nuvola apps kaboodle.svg The Secret Life of Plankton - YouTube

As well as representing the lower levels of a food chain that supports commercially important fisheries, plankton ecosystems play a role in the biogeochemical cycles of many important chemical elements, including the ocean's carbon cycle. [52] Fish larvae mainly eat zooplankton, which in turn eat phytoplankton [53]

Carbon cycle

Primarily by grazing on phytoplankton, zooplankton provide carbon to the planktic foodweb, either respiring it to provide metabolic energy, or upon death as biomass or detritus. Organic material tends to be denser than seawater, so it sinks into open ocean ecosystems away from the coastlines, transporting carbon along with it. This process, called the biological pump, is one reason that oceans constitute the largest carbon sink on Earth. However, it has been shown to be influenced by increments of temperature. [54] [55] [56] [57] In 2019, a study indicated that at ongoing rates of seawater acidification, Antarctic phytoplanktons could become smaller and less effective at storing carbon before the end of the century. [58]

It might be possible to increase the ocean's uptake of carbon dioxide (CO
2
) generated through human activities by increasing plankton production through iron fertilization – introducing amounts of iron into the ocean. However, this technique may not be practical at a large scale. Ocean oxygen depletion and resultant methane production (caused by the excess production remineralising at depth) is one potential drawback. [59] [60]

Oxygen production

Phytoplankton absorb energy from the Sun and nutrients from the water to produce their own nourishment or energy. In the process of photosynthesis, phytoplankton release molecular oxygen (O
2
) into the water as a waste byproduct. It is estimated that about 50% of the world's oxygen is produced via phytoplankton photosynthesis. [61] The rest is produced via photosynthesis on land by plants. [61] Furthermore, phytoplankton photosynthesis has controlled the atmospheric CO
2
/O
2
balance since the early Precambrian Eon. [62]

Absorption efficiency

The absorption efficiency (AE) of plankton is the proportion of food absorbed by the plankton that determines how available the consumed organic materials are in meeting the required physiological demands. [63] Depending on the feeding rate and prey composition, variations in absorption efficiency 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 absorption efficiency and small, dense pellets, while high feeding rates typically lead to low absorption efficiency and larger pellets with more organic content. Another contributing factor to dissolved organic matter (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. [64] [65] 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. [66]

Biomass variability

The growth of phytoplankton populations is dependent on light levels and nutrient availability. The chief factor limiting growth varies from region to region in the world's oceans. On a broad scale, growth of phytoplankton in the oligotrophic tropical and subtropical gyres is generally limited by nutrient supply, while light often limits phytoplankton growth in subarctic gyres. Environmental variability at multiple scales influences the nutrient and light available for phytoplankton, and as these organisms form the base of the marine food web, this variability in phytoplankton growth influences higher trophic levels. For example, at interannual scales phytoplankton levels temporarily plummet during El Niño periods, influencing populations of zooplankton, fishes, sea birds, and marine mammals.

The effects of anthropogenic warming on the global population of phytoplankton is an area of active research. Changes in the vertical stratification of the water column, the rate of temperature-dependent biological reactions, and the atmospheric supply of nutrients are expected to have important impacts on future phytoplankton productivity. [67] Additionally, changes in the mortality of phytoplankton due to rates of zooplankton grazing may be significant.

Marine phytoplankton cycling throughout water column Cycling of marine phytoplankton.png
Marine phytoplankton cycling throughout water column
Amphipod with curved exoskeleton and two long and two short antennae Amphipodredkils.jpg
Amphipod with curved exoskeleton and two long and two short antennae

Plankton diversity

Planktonic relationships

Fish and plankton

Zooplankton are the initial prey item for almost all fish larvae as they switch from their yolk sacs to external feeding. Fish rely on the density and distribution of zooplankton to match that of new larvae, which can otherwise starve. Natural factors (e.g., current variations, temperature changes) and man-made factors (e.g. river dams, ocean acidification, rising temperatures) can strongly affect zooplankton populations, which can in turn strongly affect fish larval survival, and therefore breeding success.

It has been shown that plankton can be patchy in marine environments where there aren't significant fish populations and additionally, where fish are abundant, zooplankton dynamics are influenced by the fish predation rate in their environment. Depending on the predation rate, they could express regular or chaotic behavior. [69]

A negative effect that fish larvae can have on planktonic algal blooms is that the larvae will prolong the blooming event by diminishing available zooplankton numbers; this in turn permits excessive phytoplankton growth allowing the bloom to flourish . [53]

The importance of both phytoplankton and zooplankton is also well-recognized in extensive and semi-intensive pond fish farming. Plankton population-based pond management strategies for fish rearing have been practiced by traditional fish farmers for decades, illustrating the importance of plankton even in man-made environments.

Whales and plankton

Of all animal fecal matter, it is whale feces that is the 'trophy' in terms of increasing nutrient availability. Phytoplankton are the powerhouse of open ocean primary production and they can acquire many nutrients from whale feces. [70] In the marine food web, phytoplankton are at the base of the food web and are consumed by zooplankton & krill, which are preyed upon by larger and larger marine organisms, including whales, so it can be said that whale poop fuels the entire food web.

Humans and plankton

Plankton have many direct and indirect effects on humans.

Around 70% of the oxygen in the atmosphere is produced in the oceans from phytoplankton performing photosynthesis, meaning that the majority of the oxygen available for us and other organisms that respire aerobically is produced by plankton. [71]

Plankton also make up the base of the marine food web, providing food for all the trophic levels above. Recent studies have analyzed the marine food web to see if the system runs on a top-down or bottom-up approach. Essentially, this research is focused on understanding whether changes in the food web are driven by nutrients at the bottom of the food web or predators at the top. The general conclusion is that the bottom-up approach seemed to be more predictive of food web behavior. [72] This indicates that plankton have more sway in determining the success of the primary consumer species that prey on them than do the secondary consumers that prey on the primary consumers.

In some cases, plankton act as an intermediate host for deadly parasites in humans. One such case is that of cholera, an infection caused by several pathogenic strains of Vibrio cholerae . These species have been shown to have a symbiotic relationship with chitinous zooplankton species like copepods. These bacteria benefit not only from the food provided by the chiton from the zooplankton, but also from the protection from acidic environments. Once the copepods have been ingested by a human host, the chitinous exterior protects the bacteria from the stomach acids in the stomach and proceed to the intestines. Once there, the bacteria bind with the surface of the small intestine and the host will start developing symptoms, including extreme diarrhea, within five days. [73]

See also

Related Research Articles

<span class="mw-page-title-main">Phytoplankton</span> Autotrophic members of the plankton ecosystem

Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.

<span class="mw-page-title-main">Zooplankton</span> Heterotrophic protistan or metazoan members of the plankton ecosystem

Zooplankton are the heterotrophic component of the planktonic community, 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.

<span class="mw-page-title-main">Benthos</span> Community of organisms that live in the benthic zone

Benthos, also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone. This community lives in or near marine or freshwater sedimentary environments, from tidal pools along the foreshore, out to the continental shelf, and then down to the abyssal depths.

<span class="mw-page-title-main">Biological pump</span> Carbon capture process in oceans

The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. In other words, it is a biologically mediated process which results in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).

The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word pelagic is derived from Ancient Greek πέλαγος (pélagos) 'open sea'. The pelagic zone can be thought of as an imaginary cylinder or water column between the surface of the sea and the bottom. Conditions in the water column change with depth: pressure increases; temperature and light decrease; salinity, oxygen, micronutrients all change. In a manner analogous to stratification in the Earth's atmosphere, the water column can be divided vertically into up to five different layers, with the number of layers depending on the depth of the water.

<span class="mw-page-title-main">Holoplankton</span>

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.

<span class="mw-page-title-main">Lake ecosystem</span> Type of ecosystem

A lake ecosystem or lacustrine ecosystem includes biotic (living) plants, animals and micro-organisms, as well as abiotic (non-living) physical and chemical interactions. Lake ecosystems are a prime example of lentic ecosystems, which include ponds, lakes and wetlands, and much of this article applies to lentic ecosystems in general. Lentic ecosystems can be compared with lotic ecosystems, which involve flowing terrestrial waters such as rivers and streams. Together, these two ecosystems are examples of freshwater ecosystems.

<span class="mw-page-title-main">Diel vertical migration</span> A pattern of daily vertical movement characteristic of many aquatic species

Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The adjective "diel" comes from Latin: diēs, lit. 'day', and refers to a 24-hour period. The migration occurs when organisms move up to the uppermost layer of the water at night and return to the bottom of the daylight zone of the oceans or to the dense, bottom layer of lakes during the day. DVM is important to the functioning of deep-sea food webs and the biologically-driven sequestration of carbon.

<span class="mw-page-title-main">Ecology of the San Francisco Estuary</span>

The San Francisco Estuary together with the Sacramento–San Joaquin River Delta represents a highly altered ecosystem. The region has been heavily re-engineered to accommodate the needs of water delivery, shipping, agriculture, and most recently, suburban development. These needs have wrought direct changes in the movement of water and the nature of the landscape, and indirect changes from the introduction of non-native species. New species have altered the architecture of the food web as surely as levees have altered the landscape of islands and channels that form the complex system known as the Delta.

<span class="mw-page-title-main">Forage fish</span> Small prey fish

Forage fish, also called prey fish or bait fish, are small pelagic fish that feed on planktons and other small aquatic organisms. They are in turn preyed upon by various predators including larger fish, seabirds and marine mammals, this making them keystone species in their aquatic ecosystems.

<span class="mw-page-title-main">Marine snow</span> Shower of organic detritus in the ocean

In the deep ocean, marine snow is a continuous shower of mostly organic detritus falling from the upper layers of the water column. It is a significant means of exporting energy from the light-rich photic zone to the aphotic zone below, which is referred to as the biological pump. Export production is the amount of organic matter produced in the ocean by primary production that is not recycled (remineralised) before it sinks into the aphotic zone. Because of the role of export production in the ocean's biological pump, it is typically measured in units of carbon. The term was coined by explorer William Beebe as observed from his bathysphere. As the origin of marine snow lies in activities within the productive photic zone, the prevalence of marine snow changes with seasonal fluctuations in photosynthetic activity and ocean currents. Marine snow can be an important food source for organisms living in the aphotic zone, particularly for organisms that live very deep in the water column.

<span class="mw-page-title-main">Bacterioplankton</span> Bacterial component of the plankton that drifts in the water column

Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλαγκτός (planktós), meaning "wandering" or "drifting", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and fresh water.

<span class="mw-page-title-main">Ecosystem of the North Pacific Subtropical Gyre</span> Major circulating ecosystem of ocean currents

The North Pacific Subtropical Gyre (NPSG) is the largest contiguous ecosystem on earth. In oceanography, a subtropical gyre is a ring-like system of ocean currents rotating clockwise in the Northern Hemisphere and counterclockwise in the Southern Hemisphere caused by the Coriolis Effect. They generally form in large open ocean areas that lie between land masses.

<span class="mw-page-title-main">Marine habitat</span> Habitat that supports marine life

A marine habitat is a habitat that supports marine life. Marine life depends in some way on the saltwater that is in the sea. A habitat is an ecological or environmental area inhabited by one or more living species. The marine environment supports many kinds of these habitats.

<span class="mw-page-title-main">Planktivore</span> Aquatic organism that feeds on planktonic food

A planktivore is an aquatic organism that feeds on planktonic food, including zooplankton and phytoplankton. Planktivorous organisms encompass a range of some of the planet's smallest to largest multicellular animals in both the present day and in the past billion years; basking sharks and copepods are just two examples of giant and microscopic organisms that feed upon plankton.

<span class="mw-page-title-main">Mycoplankton</span> Fungal members of the plankton communities of aquatic ecosystems

Mycoplankton are saprotrophic members of the plankton communities of marine and freshwater ecosystems. They are composed of filamentous free-living fungi and yeasts that are associated with planktonic particles or phytoplankton. Similar to bacterioplankton, these aquatic fungi play a significant role in heterotrophicmineralization and nutrient cycling. Mycoplankton can be up to 20 mm in diameter and over 50 mm in length.

<span class="mw-page-title-main">Particulate organic matter</span>

Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 millimeters (53 μm) to 2 millimeters.

<span class="mw-page-title-main">Lipid pump</span>

The lipid pump sequesters carbon from the ocean's surface to deeper waters via lipids associated with overwintering vertically migratory zooplankton. Lipids are a class of hydrocarbon rich, nitrogen and phosphorus deficient compounds essential for cellular structures. This lipid carbon enters the deep ocean as carbon dioxide produced by respiration of lipid reserves and as organic matter from the mortality of zooplankton.

<span class="mw-page-title-main">Marine food web</span> Marine consumer-resource system

A marine food web is a food web of marine life. At the base of the ocean food web are single-celled algae and other plant-like organisms known as phytoplankton. The second trophic level is occupied by zooplankton which feed off the phytoplankton. Higher order consumers complete the web. There has been increasing recognition in recent years that marine microorganisms.

<span class="mw-page-title-main">Benthic-pelagic coupling</span> Processes that connect the benthic and pelagic zones of a body of water

Benthic-pelagic coupling are processes that connect the benthic zone and the pelagic zone through the exchange of energy, mass, or nutrients. These processes play a prominent role in both freshwater and marine ecosystems and are influenced by a number of chemical, biological, and physical forces that are crucial to functions from nutrient cycling to energy transfer in food webs.

References

  1. Lalli, C.; Parsons, T. (1993). Biological Oceanography: An Introduction. Butterworth-Heinemann. ISBN   0-7506-3384-0.
  2. Smith, David J. (July 2013). "Aeroplankton and the Need for a Global Monitoring Network". BioScience. 63 (7): 515–516. doi: 10.1525/bio.2013.63.7.3 . S2CID   86371218.
  3. "plankter". American Heritage Dictionary. Houghton Mifflin Harcourt Publishing Company. Archived from the original on 9 November 2018. Retrieved 9 November 2018.
  4. Lawton, Graham (10 February 2024). "Fungi ahoy!". New Scientist. 261 (3477): 37–39. Bibcode:2024NewSc.261b..37L. doi:10.1016/S0262-4079(24)00274-4.
  5. Dolan, John (November 2012). "Microzooplankton: the microscopic (micro) animals (zoo) of the plankton" (PDF). Institut océanographique. Archived from the original (PDF) on 4 March 2016. Retrieved 16 January 2014.
  6. Hansen, Victor (1887). "Uber die Bestimmung des Plankton's oder des im Meere treibenden Materials an Pflanzen und Thieren" [On the determination of the plankton or the material floating in the sea on plants and animals]. Fünfter Bericht der Kommission zur Wissenschaftlichen Untersuchung der Deutschen Meere (in German). 12 (12–16). Berlin, Germany: Paul Parey: 1-108 via Biodiversity Heritage Library.
  7. Karleskint, George; Turner, Richard; Small, James (2013). "17: The Open Sea". Introduction to Marine Biology (4th ed.). Brooks/Cole. pp. 442–443. ISBN   978-1-133-36446-7.
  8. Agrawai, Anju; Gopnal, Krishna (2013). Biomonitoring of Water and Waste Water. Springer India. p. 34. ISBN   978-8-132-20864-8 . Retrieved 2 April 2018.
  9. "Plankter – marine biology". Encyclopædia Britannica.
  10. Emiliani, C. (1991). "Planktic/Planktonic, Nektic/Nektonic, Benthic/Benthonic". Journal of Paleontology. 65 (2): 329. Bibcode:1991JPal...65..329E. doi:10.1017/S0022336000020576. JSTOR   1305769. S2CID   131283465.
  11. Wang, G.; Wang, X.; Liu, X.; Li, Q. (2012). "Diversity and biogeochemical function of planktonic fungi in the ocean". In Raghukumar, Chandralata (ed.). Biology of Marine Fungi. Springer Berlin Heidelberg. pp. 71–88. ISBN   978-3-642-23342-5.
  12. Wommack, K.E.; Colwell, R.R. (March 2000). "Virioplankton: viruses in aquatic ecosystems". Microbiology and Molecular Biology Reviews. 64 (1): 69–114. doi:10.1128/MMBR.64.1.69-114.2000. PMC   98987 . PMID   10704475.
  13. "Plankton". Resource Library. National Geographic . Retrieved 13 September 2019.
  14. Leles, Suzana Gonçalves (November 2018). "Modelling mixotrophic functional diversity and implications for ecosystem function - Oxford Journals". Journal of Plankton Research. 40 (6): 627–642. doi:10.1093/plankt/fby044.
  15. Hartmann, M.; Grob, C.; Tarran, G.A.; Martin, A.P.; Burkill, P.H.; Scanlan, D.J.; Zubkov, M.V. (2012). "Mixotrophic basis of Atlantic oligotrophic ecosystems". Proc. Natl. Acad. Sci. USA. 109 (15): 5756–5760. Bibcode:2012PNAS..109.5756H. doi: 10.1073/pnas.1118179109 . PMC   3326507 . PMID   22451938.
  16. Ward, B.A.; Follows, M.J. (2016). "Marine mixotrophy increases trophic transfer efficiency, mean organism size, and vertical carbon flux". Proc. Natl. Acad. Sci. USA. 113 (11): 2958–2963. Bibcode:2016PNAS..113.2958W. doi: 10.1073/pnas.1517118113 . PMC   4801304 . PMID   26831076.
  17. "Mixing It Up in the Web of Life". The Scientist Magazine.
  18. "Uncovered: the mysterious killer triffids that dominate life in our oceans". 3 November 2016.
  19. "Catastrophic Darkness". Astrobiology Magazine. Archived from the original on 2015-09-26. Retrieved 2019-11-27.
  20. Chust, G., Vogt, M., Benedetti, F., Nakov, T., Villéger, S., Aubert, A., Vallina, S.M., Righetti, D., Not, F., Biard, T. and Bittner, L.(2017) "Mare incognitum: A glimpse into future plankton diversity and ecology research". Frontiers in Marine Science, 4: 68. doi : 10.3389/fmars.2017.00068.
  21. Omori, M.; Ikeda, T. (1992). Methods in Marine Zooplankton Ecology. Malabar, USA: Krieger Publishing Company. ISBN   978-0-89464-653-9.
  22. Dusenbery, David B. (2009). Living at micro scale: the unexpected physics of being small. Cambridge: Harvard University Press. ISBN   978-0-674-03116-6.
  23. Karsenti, Eric; Acinas, Silvia G.; Bork, Peer; Bowler, Chris; De Vargas, Colomban; Raes, Jeroen; Sullivan, Matthew; Arendt, Detlev; Benzoni, Francesca; Claverie, Jean-Michel; Follows, Mick; Gorsky, Gaby; Hingamp, Pascal; Iudicone, Daniele; Jaillon, Olivier; Kandels-Lewis, Stefanie; Krzic, Uros; Not, Fabrice; Ogata, Hiroyuki; Pesant, Stéphane; Reynaud, Emmanuel Georges; Sardet, Christian; Sieracki, Michael E.; Speich, Sabrina; Velayoudon, Didier; Weissenbach, Jean; Wincker, Patrick (2011). "A Holistic Approach to Marine Eco-Systems Biology". PLOS Biology. 9 (10): e1001177. doi: 10.1371/journal.pbio.1001177 . PMC   3196472 . PMID   22028628.
  24. A. C. Hardy and P. S. Milne (1938) Studies in the Distribution of Insects by Aerial Currents. Journal of Animal Ecology, 7(2):199-229
  25. Liss, P. S. (1997). The sea surface and global change. Cambridge New York: Cambridge University Press. ISBN   978-0-521-56273-7. OCLC   34933503.
  26. 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.
  27. Wallace Jr., G.T., Duce, R.A., 1978. Transport of particulate organic matter by bubbles in marine waters. Limnol. Oceanogr. 23 Ž6., 1155–1167.
  28. 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.
  29. Klassen, R. D., & Roberge, P. R. (1999). Aerosol transport modeling as an aid to understanding atmospheric corrosivity patterns. Materials & Design, 20, 159–168.
  30. 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.
  31. Chow, J. C., Watson, J. G., Green, M. C., Lowenthal, D. H., Bates, B., Oslund, W., & Torre, G. (2000). Cross-border transport and spatial variability of suspended particles in Mexicali and California's Imperial Valley. Atmospheric Environment, 34, 1833–1843.
  32. 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.
  33. 1 2 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.
  34. 1 2 3 Stübner, E. I.; Søreide, J. E. (2016-01-27). "Year-round meroplankton dynamics in high-Arctic Svalbard". Journal of Plankton Research. 38 (3): 522–536. doi: 10.1093/plankt/fbv124 .
  35. Hays, Graeme C.; Doyle, Thomas K.; Houghton, Jonathan D.R. (2018). "A Paradigm Shift in the Trophic Importance of Jellyfish?". Trends in Ecology & Evolution. 33 (11): 874–884. Bibcode:2018TEcoE..33..874H. doi:10.1016/j.tree.2018.09.001. PMID   30245075. S2CID   52336522.
  36. Lalli, C.M. & Parsons, T.R. (2001) Biological Oceanography. Butterworth-Heinemann.
  37. Johnsen, S. (2000) Transparent Animals. Scientific American282: 62-71.
  38. Nouvian, C. (2007) The Deep. University of Chicago Press.
  39. What are Ichthyoplankton? Southwest Fisheries Science Center, NOAA. Modified 3 September 2007. Retrieved 22 July 2011.
  40. Allen, Dr. Larry G.; Horn, Dr. Michael H. (15 February 2005). The Ecology of Marine Fishes: California and Adjacent Waters. University of California Press. pp. 269–319. ISBN   9780520932470.
  41. Harvey, Edmund Newton (1952). Bioluminescence. Academic Press.
  42. Anderson, Genny. "Marine Plankton". Marine Science. Retrieved 2012-04-04.
  43. Talks, Ted. "Zooplankton". Marine Life/Marine Invertebrates. Archived from the original on 2017-12-07. Retrieved 2012-04-04.
  44. "Plankton". Britannica. Retrieved 2020-06-13.
  45. Ershova, E. A.; Descoteaux, R. (2019-08-13). "Diversity and Distribution of Meroplanktonic Larvae in the Pacific Arctic and Connectivity With Adult Benthic Invertebrate Communities". Frontiers in Marine Science. 6. doi: 10.3389/fmars.2019.00490 . hdl: 10037/16483 . S2CID   199638114.
  46. Sorokin, Yuri I. (12 March 2013). Coral Reef Ecology. Springer Science & Business Media. p. 96. ISBN   9783642800467.
  47. 1 2 Chapman, Michael J.; Margulis, Lynn (2009). Kingdoms and Domains: An Illustrated Guide to the Phyla of Life on Earth ([4th ed.]. ed.). Amsterdam: Academic Press/Elsevier. pp.  566. ISBN   978-0123736215.
  48. Simberloff, Daniel; Rejmanek, Marcel, eds. (2011). Encyclopedia of biological invasions . Berkeley: University of California Press. pp.  736. ISBN   978-0520264212.
  49. Kennish, Michael J., ed. (2004). Estuarine Research, Monitoring, and Resource Protection. Boca Raton, Fla.: CRC Press. p. 194. ISBN   978-0849319600. Archived from the original on 2013-01-20.
  50. Martin, J.H.; Fitzwater, S.E. (1988). "Iron-deficiency limits phytoplankton growth in the Northeast Pacific Subarctic". Nature. 331 (6154): 341–343. Bibcode:1988Natur.331..341M. doi:10.1038/331341a0. S2CID   4325562.
  51. Boyd, P.W.; et al. (2000). "A mesoscale phytoplankton bloom in the polar Southern Ocean stimulated by fertilization". Nature. 407 (6805): 695–702. Bibcode:2000Natur.407..695B. doi:10.1038/35037500. PMID   11048709. S2CID   4368261.
  52. Falkowski, Paul G. (1994). "The role of phytoplankton photosynthesis in global biogeochemical cycles" (PDF). Photosynthesis Research. 39 (3): 235–258. Bibcode:1994PhoRe..39..235F. doi:10.1007/BF00014586. PMID   24311124. S2CID   12129871.[ permanent dead link ]
  53. 1 2 James, Alex; Pitchford, Jonathan W.; Brindley, John (2003-02-01). "The relationship between plankton blooms, the hatching of fish larvae, and recruitment". Ecological Modelling. 160 (1): 77–90. Bibcode:2003EcMod.160...77J. doi:10.1016/S0304-3800(02)00311-3. ISSN   0304-3800.
  54. Sarmento, H.; Montoya, JM.; Vázquez-Domínguez, E.; Vaqué, D.; Gasol, JM. (2010). "Warming effects on marine microbial food web processes: how far can we go when it comes to predictions?". Philosophical Transactions of the Royal Society B: Biological Sciences. 365 (1549): 2137–2149. doi:10.1098/rstb.2010.0045. PMC   2880134 . PMID   20513721.
  55. Vázquez-Domínguez, E.; Vaqué, D.; Gasol, JM. (2007). "Ocean warming enhances respiration and carbon demand of coastal microbial plankton". Global Change Biology. 13 (7): 1327–1334. Bibcode:2007GCBio..13.1327V. doi:10.1111/j.1365-2486.2007.01377.x. hdl: 10261/15731 . S2CID   8721854.
  56. Vázquez-Domínguez, E.; Vaqué, D.; Gasol, JM. (2012). "Temperature effects on the heterotrophic bacteria, heterotrophic nanoflagellates, and microbial top predators of NW Mediterranean". Aquatic Microbial Ecology. 67 (2): 107–121. doi: 10.3354/ame01583 . hdl: 10261/95626 .
  57. Mazuecos, E.; Arístegui, J.; Vázquez-Domínguez, E.; Ortega-Retuerta, E.; Gasol, JM.; Reche, I. (2012). "Temperature control of microbial respiration and growth efficiency in the mesopelagic zone of the South Atlantic and Indian Oceans". Deep Sea Research Part I: Oceanographic Research Papers. 95 (2): 131–138. doi: 10.3354/ame01583 . hdl: 10261/95626 .
  58. Petrou, Katherina; Nielsen, Daniel (2019-08-27). "Acid oceans are shrinking plankton, fueling faster climate change". phys.org. Retrieved 2019-09-07.
  59. Chisholm, S.W.; et al. (2001). "Dis-crediting ocean fertilization". Science. 294 (5541): 309–310. doi:10.1126/science.1065349. PMID   11598285. S2CID   130687109.
  60. Aumont, O.; Bopp, L. (2006). "Globalizing results from ocean in situ iron fertilization studies". Global Biogeochemical Cycles. 20 (2): GB2017. Bibcode:2006GBioC..20.2017A. doi: 10.1029/2005GB002591 .
  61. 1 2 Roach, John (June 7, 2004). "Source of Half Earth's Oxygen Gets Little Credit". National Geographic News. Archived from the original on June 8, 2004. Retrieved 2016-04-04.
  62. Tappan, Helen (April 1968). "Primary production, isotopes, extinctions and the atmosphere". Palaeogeography, Palaeoclimatology, Palaeoecology. 4 (3): 187–210. Bibcode:1968PPP.....4..187T. doi:10.1016/0031-0182(68)90047-3.
  63. Steinberg, Deborah K.; Landry, Michael R. (2017). "Zooplankton and the Ocean Carbon Cycle". Annual Review of Marine Science. 9: 413–444. Bibcode:2017ARMS....9..413S. doi:10.1146/annurev-marine-010814-015924. PMID   27814033.
  64. Moller, E. F. (2004). "Sloppy feeding in marine copepods: Prey-size-dependent production of dissolved organic carbon". Journal of Plankton Research. 27: 27–35. doi: 10.1093/plankt/fbh147 .
  65. Møller, Eva Friis (2007). "Production of dissolved organic carbon by sloppy feeding in the copepods Acartia tonsa, Centropages typicus, and Temora longicornis". Limnology and Oceanography. 52 (1): 79–84. Bibcode:2007LimOc..52...79M. doi: 10.4319/lo.2007.52.1.0079 .
  66. Thor, P.; Dam, HG; Rogers, DR (2003). "Fate of organic carbon released from decomposing copepod fecal pellets in relation to bacterial production and ectoenzymatic activity". Aquatic Microbial Ecology. 33: 279–288. doi: 10.3354/ame033279 .
  67. Steinacher, M.; et al. (2010). "Projected 21st century decrease in marine productivity: a multi-model analysis". Biogeosciences. 7 (3): 979–1005. Bibcode:2010BGeo....7..979S. doi: 10.5194/bg-7-979-2010 . hdl: 11858/00-001M-0000-0011-F69E-5 .
  68. Michael Le Page (March 2019). "Animal with an anus that comes and goes could reveal how ours evolved". New Scientist.
  69. Medvinsky, Alexander B.; Tikhonova, Irene A.; Aliev, Rubin R.; Li, Bai-Lian; Lin, Zhen-Shan; Malchow, Horst (2001-07-26). "Patchy environment as a factor of complex plankton dynamics". Physical Review E. 64 (2): 021915. Bibcode:2001PhRvE..64b1915M. doi:10.1103/PhysRevE.64.021915. ISSN   1063-651X. PMID   11497628.
  70. "whale poop and phytoplankton, fighting climate change". IFAW. Retrieved 2022-03-29.
  71. Sekerci, Yadigar; Petrovskii, Sergei (2015-12-01). "Mathematical Modelling of Plankton–Oxygen Dynamics Under the Climate Change". Bulletin of Mathematical Biology. 77 (12): 2325–2353. doi:10.1007/s11538-015-0126-0. hdl: 2381/36058 . ISSN   1522-9602. PMID   26607949. S2CID   8637912.
  72. Frederiksen, Morten; Edwards, Martin; Richardson, Anthony J.; Halliday, Nicholas C.; Wanless, Sarah (November 2006). "From plankton to top predators: bottom-up control of a marine food web across four trophic levels". Journal of Animal Ecology. 75 (6): 1259–1268. Bibcode:2006JAnEc..75.1259F. doi:10.1111/j.1365-2656.2006.01148.x. ISSN   0021-8790. PMID   17032358.
  73. Lipp, Erin K.; Huq, Anwar; Colwell, Rita R. (October 2002). "Effects of Global Climate on Infectious Disease: the Cholera Model". Clinical Microbiology Reviews. 15 (4): 757–770. doi:10.1128/CMR.15.4.757-770.2002. ISSN   0893-8512. PMC   126864 . PMID   12364378.

Further reading