Paradox of the plankton

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.

Diatoms through the microscope
Marine diatoms are among the many planktonic organisms that paradoxically appear to flout the competitive exclusion principle.

Ecological paradox

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. 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.[3] More recent work has proposed that the paradox can be resolved by factors such as: chaotic fluid motion;[4] size-selective grazing;[5] spatio-temporal heterogeneity;[6] and environmental fluctuations.[7] More generally, some 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.[8] In Mitchell et al. (2008), researchers found that small-scale analysis of plankton distribution exhibited patches of aggregation, on the order of 10 cm, that had sufficient lifetimes (> 10 minutes) to enable plankton grazing, competition, and infection.[9]

See also


  1. ^ Hardin, G. (1960). "The Competitive Exclusion Principle". Science. 131 (3409): 1292–1297. doi:10.1126/science.131.3409.1292. PMID 14399717.
  2. ^ Gause, G. F. (1932). "Experimental Studies on the Struggle for Existence - I. Mixed Population of Two Species of Yeast". Journal of Experimental Biology. 9: 389–402.
  3. ^ Hutchinson, G. E. (1961) The paradox of the plankton. American Naturalist 95, 137-145.
  4. ^ Károlyi, G., Péntek, Á., Scheuring, I., Tél, T., Toroczkai, Z. (2000) Chaotic flow: the physics of species coexistence. Proceedings of the National Academy of Sciences 97, 13661-13665.
  5. ^ Wiggert, J.D., Haskell, A.G.E., Paffenhofer, G.A., Hofmann, E.E. and Klinck, J.M. (2005) The role of feeding behavior in sustaining copepod populations in the tropical ocean Archived 2008-09-05 at the Wayback Machine. Journal of Plankton Research 27, 1013-1031.
  6. ^ Miyazaki, T., Tainaka, K., Togashi, T., Suzuki, T. and Yoshimura, J. (2006) Spatial coexistence of phytoplankton species in ecological timescale Archived 2007-09-27 at the Wayback Machine. Population Ecology 48(2), 107-112.
  7. ^ Descamps-Julien, B.; Gonzalez, A. (2005). "Stable coexistence in a fluctuating environment: An experimental demonstration" (PDF). Ecology. 86 (10): 2815–2824. doi:10.1890/04-1700. Archived from the original (PDF) on 17 November 2006. Retrieved 18 October 2014.
  8. ^ Scheffer, M., Rinaldi, S., Huisman, J. and Weissing, F.J. (2003) Why plankton communities have no equilibrium: solutions to the paradox. Hydrobiologia 491, 9-18.
  9. ^ Mitchell, J.G., Yamazaki, H., Seuront, L., Wolk, F., Li, H. (2008) Phytoplankton patch patterns: Seascape anatomy in a turbulent ocean. Journal of Marine Systems 69, 247-253.

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Aquatic ecosystem

An aquatic ecosystem is an ecosystem in a body of water. Communities of organisms that are dependent on each other and on their environment live in aquatic ecosystems. The two main types of aquatic ecosystems are marine ecosystems and freshwater ecosystems.


Benthos is the community of organisms that live on, in, or near the seabed, river, lake, or stream bottom, 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.

Many organisms adapted to deep-water pressure cannot survive in the upperparts of the water column. The pressure difference can be very significant (approximately one atmosphere for each 10 metres of water depth).Because light is absorbed before it can reach deep ocean-water, the energy source for deep benthic ecosystems is often organic matter from higher up in the water column that drifts down to the depths. This dead and decaying matter sustains the benthic food chain; most organisms in the benthic zone are scavengers or detritivores.

The term benthos, coined by Haeckel in 1891, comes from the Greek noun βένθος "depth of the sea". Benthos is used in freshwater biology to refer to organisms at the bottom of freshwater bodies of water, such as lakes, rivers, and streams. There is also a redundant synonym, benthon.

Cascade effect (ecology)

An ecological cascade effect is a series of secondary extinctions that is triggered by the primary extinction of a key species in an ecosystem. Secondary extinctions are likely to occur when the threatened species are: dependent on a few specific food sources, mutualistic (dependent on the key species in some way), or forced to coexist with an invasive species that is introduced to the ecosystem. Species introductions to a foreign ecosystem can often devastate entire communities, and even entire ecosystems. These exotic species monopolize the ecosystem's resources, and since they have no natural predators to decrease their growth, they are able to increase indefinitely. Olsen et al. showed that exotic species have caused lake and estuary ecosystems to go through cascade effects due to loss of algae, crayfish, mollusks, fish, amphibians, and birds. However, the principal cause of cascade effects is the loss of top predators as the key species. As a result of this loss, a dramatic increase (ecological release) of prey species occurs. The prey is then able to overexploit its own food resources, until the population numbers decrease in abundance, which can lead to extinction. When the prey's food resources disappear, they starve and may go extinct as well. If the prey species is herbivorous, then their initial release and exploitation of the plants may result in a loss of plant biodiversity in the area. If other organisms in the ecosystem also depend upon these plants as food resources, then these species may go extinct as well. An example of the cascade effect caused by the loss of a top predator is apparent in tropical forests. When hunters cause local extinctions of top predators, the predators' prey's population numbers increase, causing an overexploitation of a food resource and a cascade effect of species loss. Recent studies have been performed on approaches to mitigate extinction cascades in food-web networks.

Competitive exclusion principle

In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition named for Georgy Gause that two species competing for the same limiting 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 can not coexist".

Floodplain restoration

Floodplain restoration is the process of fully or partially restoring a river's floodplain to its original conditions before having been affected by the construction of levees (dikes) and the draining of wetlands and marshes.

The objectives of restoring floodplains include the reduction of the incidence of floods, the provision of habitats for aquatic species, the improvement of water quality and the increased recharge of groundwater.

G. Evelyn Hutchinson

George Evelyn Hutchinson (January 30, 1903 – May 17, 1991), was a British ecologist sometimes described as the "father of modern ecology." He contributed for more than sixty years to the fields of limnology, systems ecology, radiation ecology, entomology, genetics, biogeochemistry, a mathematical theory of population growth, art history, philosophy, religion, and anthropology. He worked on the passage of phosphorus through lakes, the chemistry and biology of lakes, the theory of interspecific competition, and on insect taxonomy and genetics, zoo-geography and African water bugs. He is known as one of the first to combine ecology with mathematics. He became an international expert on lakes and wrote the four-volume Treatise on Limnology in 1957.Hutchinson earned his degree in zoology from Cambridge University but chose not to earn a doctorate, of which he came to be proud as he aged. Although born in England, he spent nearly his entire professional life at Yale University in the United States where he was Sterling Professor of Zoology and focused on working with graduate students.


Limnology ( lim-NOL-ə-jee; from Greek λίμνη, limne, "lake" and λόγος, logos, "knowledge"), is the study of inland aquatic ecosystems.

The study of limnology includes aspects of the biological, chemical, physical, and geological characteristics and functions of inland waters (running and standing waters, fresh and saline, natural or man-made). This includes the study of lakes, reservoirs, ponds, rivers, springs, streams, wetlands, and groundwater. A more recent sub-discipline of limnology, termed landscape limnology, studies, manages, and seeks to conserve these ecosystems using a landscape perspective, by explicitly examining connections between an aquatic ecosystem and its watershed. Recently, the need to understand global inland waters as part of the Earth System created a sub-discipline called global limnology. This approach considers processes in inland waters on a global scale, like the role of inland aquatic ecosystems in global biogeochemical cycles.Limnology is closely related to aquatic ecology and hydrobiology, which study aquatic organisms and their interactions with the abiotic (non-living) environment. While limnology has substantial overlap with freshwater-focused disciplines (e.g., freshwater biology), it also includes the study of inland salt lakes.

List of unsolved problems in biology

This article lists currently unsolved problems in biology.

List of watershed topics

This list embraces topographical watersheds and drainage basins and other topics focused on them.

Marine bacteriophage

Marine bacteriophages or marine phages are viruses that live as obligate parasitic agents in marine bacteria such as cyanobacteria. Their existence was discovered through electron microscopy and epifluorescence microscopy of ecological water samples, and later through metagenomic sampling of uncultured viral samples. Marine phages, although microscopic and essentially unnoticed by scientists until recently, appear to be the most abundant and diverse form of DNA replicating agent on the planet. There are approximately 4x1030 phage in oceans or 5x107 per millilitre. Quantification of marine viruses was originally performed using transmission electron microscopy but has been replaced by epifluorescence or flow cytometry.

Mesopredator release hypothesis

The mesopredator release hypothesis is an ecological theory used to describe the interrelated population dynamics between apex predators and mesopredators within an ecosystem, such that a collapsing population of the former results in dramatically-increased populations of the latter. This hypothesis describes the phenomenon of trophic cascade in specific terrestrial communities.

A mesopredator is a medium-sized, middle trophic level predator, which both preys and is preyed upon. Examples are raccoons, skunks, snakes, cownose rays, and small sharks.

Particle (ecology)

In marine and freshwater ecology, a particle is a small object. Particles can remain in suspension in the ocean or freshwater. However, they eventually settle (rate determined by Stokes' law) and accumulate as sediment. Some can enter the atmosphere through wave action where they can act as cloud condensation nuclei (CCN). Many organisms filter particles out of the water with unique filtration mechanisms (filter feeders). Particles are often associated with high loads of toxins which attach to the surface. As these toxins are passed up the food chain they accumulate in fatty tissue and become increasingly concentrated in predators (see bioaccumulation). Very little is known about the dynamics of particles, especially when they are re-suspended by dredging. They can remain floating in the water and drift over long distances. The decomposition of some particles by bacteria consumes a lot of oxygen and can cause the water to become hypoxic.

Photic zone

The photic zone, euphotic zone (Greek for "well lit": εὖ "well" + φῶς "light"), or sunlight (or sunlit) zone is the uppermost layer of water in a lake or ocean that is exposed to intense sunlight. It corresponds roughly to the layer above the compensation point, i.e. depth where the rate of carbon dioxide uptake, or equivalently, the rate of photosynthetic oxygen production, is equal to the rate of carbon dioxide production, equivalent to the rate of respiratory oxygen consumption, i.e. the depth where net carbon dioxide assimilation is zero.

It extends from the surface down to a depth where light intensity falls to one percent of that at the surface, called the euphotic depth. Accordingly, its thickness depends on the extent of light attenuation in the water column. Typical euphotic depths vary from only a few centimetres in highly turbid eutrophic lakes, to around 200 meters in the open ocean. It also varies with seasonal changes in turbidity.

Since the photic zone is where almost all of the photosynthesis occurs, the depth of the photic zone is generally proportional to the level of primary production that occurs in that area of the ocean. About 90% of all marine life lives in the photic zone. A small amount of primary production is generated deep in the abyssal zone around the hydrothermal vents which exist along some mid-oceanic ridges.

The zone which extends from the base of the euphotic zone to about 200 metres is sometimes called the disphotic zone. While there is some light, it is insufficient for photosynthesis, or at least insufficient for photosynthesis at a rate greater than respiration. The euphotic zone together with the disphotic zone coincides with the epipelagic zone. The bottommost zone, below the euphotic zone, is called the aphotic zone. Most deep ocean waters belong to this zone.

The transparency of the water, which determines the depth of the photic zone, is measured simply with a Secchi disk. It may also be measured with a photometer lowered into the water.


Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of oceans, seas and freshwater basin ecosystems. The name comes from the Greek words φυτόν (phyton), meaning "plant", and πλανκτός (planktos), meaning "wanderer" or "drifter". Most phytoplankton are too small to be individually seen with the unaided eye. However, when present in high enough numbers, some varieties may be noticeable as colored patches on the water surface due to the presence of chlorophyll within their cells and accessory pigments (such as phycobiliproteins or xanthophylls) in some species.


Plankton are the diverse collection of organisms that live in large bodies of water and are unable to swim against a current. The individual organisms constituting plankton are called plankters. They provide a crucial source of food to many large aquatic organisms, such as fish and whales.

These organisms include bacteria, archaea, algae, protozoa and drifting or floating animals that inhabit—for example—the pelagic zone of oceans, seas, or bodies of fresh water. Essentially, plankton are defined by their ecological niche rather than any phylogenetic or taxonomic classification.

Though many planktonic species are microscopic in size, plankton includes organisms over a wide range of sizes, including large organisms such as jellyfish.

Technically the term does not include organisms on the surface of the water, which are called pleuston—or those that swim actively in the water, which are called nekton.

Productivity (ecology)

In ecology, productivity refers to the rate of generation of biomass in an ecosystem. It is usually expressed in units of mass per unit surface (or volume) per unit time, for instance grams per square metre per day (g m−2 d−1). The mass unit may relate to dry matter or to the mass of carbon generated. Productivity of autotrophs such as plants is called primary productivity, while that of heterotrophs such as animals is called secondary productivity.

Ramsar site

A Ramsar site is a wetland site designated to be of international importance under the Ramsar Convention.The Convention on Wetlands, known as the Ramsar Convention, is an intergovernmental environmental treaty established in 1971 by UNESCO, which came into force in 1975. It provides for national action and international cooperation regarding the conservation of wetlands, and wise sustainable use of their resources.Ramsar identifies wetlands of international importance, especially those providing waterfowl habitat.

As of 2016, there were 2,231 Ramsar sites, protecting 214,936,005 hectares (531,118,440 acres), and 169 national governments are currently participating.

Sustainable gardening

Sustainable gardening includes the more specific sustainable landscapes, sustainable landscape design, sustainable landscaping, sustainable landscape architecture, resulting in sustainable sites. It comprises a disparate group of horticultural interests that can share the aims and objectives associated with the international post-1980s sustainable development and sustainability programs developed to address the fact that humans are now using natural biophysical resources faster than they can be replenished by nature.Included within this compass are those home gardeners, and members of the landscape and nursery industries, and municipal authorities, that integrate environmental, social, and economic factors to create a more sustainable future.

Organic gardening and the use of native plants are integral to sustainable gardening.


Zooplankton (, ) 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". Individual zooplankton are usually microscopic, but some (such as jellyfish) are larger and visible to the naked eye.

Patterns of evolution
About plankton
By size
Related topics
Aquatic ecosystems
Food webs
Example webs
Ecology: Modelling ecosystems: Other components

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