In aquatic biology, the paradox of the plankton describes the situation in which a limited range of resources supports an unexpectedly wide range of plankton species, apparently flouting the competitive exclusion principle, which holds that when two species compete for the same resource, one will be driven to extinction.
The paradox of the plankton results from the clash between the observed diversity of plankton and the competitive exclusion principle, [1] also known as Gause's law, [2] which states that, when two species compete for the same resource, ultimately only one will persist and the other will be driven to extinction. Coexistence between two such species is impossible because the dominant one will inevitably deplete the shared resources, thus decimating the inferior population. [3] Phytoplankton life is diverse at all phylogenetic levels despite the limited range of resources (e.g. light, nitrate, phosphate, silicic acid, iron) for which they compete amongst themselves. The paradox of the plankton was originally described in 1961 by G. Evelyn Hutchinson, who proposed that the paradox could be resolved by factors such as vertical gradients of light or turbulence, symbiosis or commensalism, differential predation, or constantly changing environmental conditions. [4]
Later studies found that the paradox can be resolved by factors such as: zooplankton grazing pressure; [5] chaotic fluid motion; [6] size-selective grazing; [7] spatio-temporal heterogeneity; [8] bacterial mediation; [9] or environmental fluctuations. [10] In general, researchers suggest that ecological and environmental factors continually interact such that the planktonic habitat never reaches an equilibrium for which a single species is favoured. [11]
While it was long assumed that turbulence disrupts plankton patches at spatial scales less than a few metres, researchers using small-scale analysis of plankton distribution found that these exhibited patches of aggregation (on the order of 10cm) that had sufficient lifetimes (more than 10 minutes) to enable plankton grazing, competition, and infection. [12]
One potential resolution to the paradox is the control on plankton populations by marine lytic viruses. Marine viruses play an important role in bacteria and plankton ecology. They are a significant component of biogeochemical cycling [13] and horizontal gene transfer in both bacterial and plankton communities. Viruses are the most abundant organisms in the ocean, and have the capacity to deplete host populations very rapidly. Marine viruses infect specific host species, and therefore an abundance of a virus can quickly and effectively alter the structure of the phytoplankton and bacterial communities. Via the lytic cycle, a virus encounters a host and reproduces until the cell bursts, releasing viruses. Viruses can also enter a lysogenic cycle, in which the virus writes its DNA into the host genome. When a phytoplankton species enters a bloom period, cell concentration increases and many viral targets suddenly become available. [14]
One explanation to the paradox of the plankton is the "Boom-and-busted dynamic" hypothesis, also called "Kill the winner." In a phytoplankton bloom, an individual species multiplies rapidly in ideal conditions, which increases its cell concentration in an area, outcompeting other phytoplankton. This "boom" in host cells creates an opportunity for rapid infection by viruses, leading to a "bust" in which the phytoplankton population rapidly diminishes. This creates a large gap in the local phytoplankton ecology and allows other species to fill in and continue growing. Such population control by viruses creates temporal and spatial diversity in phytoplankton communities. Long term control results, as the virus prevents the formerly dominant species from booming during future bloom events. [15]
Plankton are the diverse collection of organisms found in water that are unable to propel themselves against a current. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish, and baleen whales.
Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdom Protista, according to Robert Whittaker's five-kingdom system, or clade Hacrobia, according to a newer biological classification system. Within the Hacrobia, the coccolithophores are in the phylum or division Haptophyta, class Prymnesiophyceae. Coccolithophores are almost exclusively marine, are photosynthetic, and exist in large numbers throughout the sunlight zone of the ocean.
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, meaning 'wanderer' or 'drifter'.
Zooplankton are the animal 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.
In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the slightest advantage over another, the one with the advantage will dominate in the long term. This leads either to the extinction of the weaker competitor or to an evolutionary or behavioral shift toward a different ecological niche. The principle has been paraphrased in the maxim "complete competitors cannot coexist".
Cyanophages are viruses that infect cyanobacteria, also known as Cyanophyta or blue-green algae. Cyanobacteria are a phylum of bacteria that obtain their energy through the process of photosynthesis. Although cyanobacteria metabolize photoautotrophically like eukaryotic plants, they have prokaryotic cell structure. Cyanophages can be found in both freshwater and marine environments. Marine and freshwater cyanophages have icosahedral heads, which contain double-stranded DNA, attached to a tail by connector proteins. The size of the head and tail vary among species of cyanophages. Cyanophages infect a wide range of cyanobacteria and are key regulators of the cyanobacterial populations in aquatic environments, and may aid in the prevention of cyanobacterial blooms in freshwater and marine ecosystems. These blooms can pose a danger to humans and other animals, particularly in eutrophic freshwater lakes. Infection by these viruses is highly prevalent in cells belonging to Synechococcus spp. in marine environments, where up to 5% of cells belonging to marine cyanobacterial cells have been reported to contain mature phage particles.
Phycodnaviridae is a family of large (100–560 kb) double-stranded DNA viruses that infect marine or freshwater eukaryotic algae. Viruses within this family have a similar morphology, with an icosahedral capsid. As of 2014, there were 33 species in this family, divided among 6 genera. This family belongs to a super-group of large viruses known as nucleocytoplasmic large DNA viruses. Evidence was published in 2014 suggesting that specific strains of Phycodnaviridae might infect humans rather than just algal species, as was previously believed. Most genera under this family enter the host cell by cell receptor endocytosis and replicate in the nucleus. Phycodnaviridae play important ecological roles by regulating the growth and productivity of their algal hosts. Algal species such Heterosigma akashiwo and the genus Chrysochromulina can form dense blooms which can be damaging to fisheries, resulting in losses in the aquaculture industry. Heterosigma akashiwo virus (HaV) has been suggested for use as a microbial agent to prevent the recurrence of toxic red tides produced by this algal species. Phycodnaviridae cause death and lysis of freshwater and marine algal species, liberating organic carbon, nitrogen and phosphorus into the water, providing nutrients for the microbial loop.
The microbial loop describes a trophic pathway where, in aquatic systems, dissolved organic carbon (DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by phytoplankton-zooplankton-nekton. In soil systems, the microbial loop refers to soil carbon. The term microbial loop was coined by Farooq Azam, Tom Fenchel et al. in 1983 to include the role played by bacteria in the carbon and nutrient cycles of the marine environment.
Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλανκτος, meaning "wanderer" or "drifter", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and freshwater.
Chrysochromulina is a genus of haptophytes. This phytoplankton is distributed globally in brackish and marine waters across approximately 60 known species. All Chrysochromulina species are phototrophic, however some have been shown to be mixotrophic, including exhibiting phagotrophy under certain environmental conditions. The cells are small, characterized by having scales, and typically observed using electron microscopy. Some species, under certain environmental conditions have been shown to produce toxic compounds that are harmful to larger marine life including fish.
Marine microorganisms are defined by their habitat as microorganisms living in a marine environment, that is, in the saltwater of a sea or ocean or the brackish water of a coastal estuary. A microorganism is any microscopic living organism or virus, that is too small to see with the unaided human eye without magnification. Microorganisms are very diverse. They can be single-celled or multicellular and include bacteria, archaea, viruses and most protozoa, as well as some fungi, algae, and animals, such as rotifers and copepods. Many macroscopic animals and plants have microscopic juvenile stages. Some microbiologists also classify viruses as microorganisms, but others consider these as non-living.
Akashiwo sanguinea is a species of marine dinoflagellates well known for forming blooms that result in red tides. The organism is unarmored (naked). Therefore, it lacks a thick cellulose wall, the theca, common in other genera of dinoflagellates. Reproduction of the phytoplankton species is primarily asexual.
Phycotoxins are complex allelopathic chemicals produced by eukaryotic and prokaryotic algal secondary metabolic pathways. More simply, these are toxic chemicals synthesized by photosynthetic organisms. These metabolites are not harmful to the producer but may be toxic to either one or many members of the marine food web. This page focuses on phycotoxins produced by marine microalgae; however, freshwater algae and macroalgae are known phycotoxin producers and may exhibit analogous ecological dynamics. In the pelagic marine food web, phytoplankton are subjected to grazing by macro- and micro-zooplankton as well as competition for nutrients with other phytoplankton species. Marine bacteria try to obtain a share of organic carbon by maintaining symbiotic, parasitic, commensal, or predatory interactions with phytoplankton. Other bacteria will degrade dead phytoplankton or consume organic carbon released by viral lysis. The production of toxins is one strategy that phytoplankton use to deal with this broad range of predators, competitors, and parasites. Smetacek suggested that "planktonic evolution is ruled by protection and not competition. The many shapes of plankton reflect defense responses to specific attack systems". Indeed, phytoplankton retain an abundance of mechanical and chemical defense mechanisms including cell walls, spines, chain/colony formation, and toxic chemical production. These morphological and physiological features have been cited as evidence for strong predatory pressure in the marine environment. However, the importance of competition is also demonstrated by the production of phycotoxins that negatively impact other phytoplankton species. Flagellates are the principle producers of phycotoxins; however, there are known toxigenic diatoms, cyanobacteria, prymnesiophytes, and raphidophytes. Because many of these allelochemicals are large and energetically expensive to produce, they are synthesized in small quantities. However, phycotoxins are known to accumulate in other organisms and can reach high concentrations during algal blooms. Additionally, as biologically active metabolites, phycotoxins may produce ecological effects at low concentrations. These effects may be subtle, but have the potential to impact the biogeographic distributions of phytoplankton and bloom dynamics.
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. Planktivory can be an important mechanism of top-down control that contributes to trophic cascades in aquatic and marine systems. There is a tremendous diversity of feeding strategies and behaviors that planktivores utilize to capture prey. Some planktivores utilize tides and currents to migrate between estuaries and coastal waters; other aquatic planktivores reside in lakes or reservoirs where diverse assemblages of plankton are present, or migrate vertically in the water column searching for prey. Planktivore populations can impact the abundance and community composition of planktonic species through their predation pressure, and planktivore migrations facilitate nutrient transport between benthic and pelagic habitats.
The "Kill the Winner" hypothesis (KtW) is an ecological model of population growth involving prokaryotes, viruses and protozoans that links trophic interactions to biogeochemistry. The model is related to the Lotka–Volterra equations. It assumes that prokaryotes adopt one of two strategies when competing for limited resources: priority is either given to population growth ("winners") or survival ("defenders"). As "winners" become more abundant and active in their environment, their contact with host-specific viruses increases, making them more susceptible to viral infection and lysis. Thus, viruses moderate the population size of "winners" and allow multiple species to coexist. Current understanding of KtW primarily stems from studies of lytic viruses and their host populations.
The viral shunt is a mechanism that prevents marine microbial particulate organic matter (POM) from migrating up trophic levels by recycling them into dissolved organic matter (DOM), which can be readily taken up by microorganisms. The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.
Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers is larger than the biomass of primary producers. This happens because the ocean's primary producers are tiny phytoplankton which grow and reproduce rapidly, so a small mass can have a fast rate of primary production. In contrast, many significant terrestrial primary producers, such as mature forests, grow and reproduce slowly, so a much larger mass is needed to achieve the same rate of primary production.
Marine viruses are defined by their habitat as viruses that are found in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Viruses are small infectious agents that can only replicate inside the living cells of a host organism, because they need the replication machinery of the host to do so. They can infect all types of life forms, from animals and plants to microorganisms, including bacteria and archaea.
Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.
Transparent exopolymer particles (TEPs) are extracellular acidic polysaccharides produced by phytoplankton and bacteria in saltwater, freshwater, and wastewater. They are incredibly abundant and play a significant role in biogeochemical cycling of carbon and other elements in water. Through this, they also play a role in the structure of food webs and trophic levels. TEP production and overall concentration has been observed to be higher in the Pacific Ocean compared to the Atlantic, and is more related to solar radiation in the Pacific. TEP concentration has been found to decrease with depth, having the highest concentration at the surface, especially associated with the SML, either by upward flux or sea surface production. Chlorophyll a has been found to be the best indicator of TEP concentration, rather than heterotrophic grazing abundance, further emphasizing the role of phytoplankton in TEP production. TEP concentration is especially enhanced by haptophyte phytoplanktonic dominance, solar radiation exposure, and close proximity to sea ice. TEPs also do not seem to show any diel cycles. High concentrations of TEPs in the surface ocean slow the sinking of solid particle aggregations, prolonging pelagic residence time. TEPs may provide an upward flux of materials such as bacteria, phytoplankton, carbon, and trace nutrients. High TEP concentrations were found under arctic sea ice, probably released by sympagic algae. TEP is efficiently recycled in the ocean, as heterotrophic grazers such as zooplankton and protists consume TEP and produce new TEP precursors to be reused, further emphasizing the importance of TEPs in marine carbon cycling. TEP abundance tends to be higher in coastal, shallow waters compared to deeper, oceanic waters. Diatom-dominated phytoplankton colonies produce larger, and stickier, TEPs, which may indicate that TEP size distribution and composition may be a useful tool in determining aggregate planktonic community structure.
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