Commensalism

Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed.[1] This is in contrast with mutualism, in which both organisms benefit from each other, amensalism, where one is harmed while the other is unaffected, and parasitism, where one benefits while the other is harmed. The commensal (the species that benefits from the association) may obtain nutrients, shelter, support, or locomotion from the host species, which is substantially unaffected. The commensal relation is often between a larger host and a smaller commensal; the host organism is unmodified, whereas the commensal species may show great structural adaptation consonant with its habits, as in the remoras that ride attached to sharks and other fishes. Both remora and pilot fish feed on the leftovers of their hosts' meals. Numerous birds perch on bodies of large mammal herbivores or feed on the insects turned up by grazing mammals.[2]

Spearfish remora
Remora are specially adapted to attach themselves to larger fish that provide locomotion and food.

Etymology

The word "commensalism" is derived from the word "commensal", meaning "eating at the same table" in human social interaction, which in turn comes through French from the Medieval Latin commensalis, meaning "sharing a table", from the prefix com-, meaning "together", and mensa, meaning "table" or "meal".[3] Commensality, at Oxford and Cambridge Universities, refers to professors eating at the same table as students (as they live in the same "college").

Pierre-Joseph van Beneden introduced the term "commensalism" in 1876.[4]

Examples of commensal relationships

The commensal pathway was traveled by animals that fed on refuse around human habitats or by animals that preyed on other animals drawn to human camps. Those animals established a commensal relationship with humans in which the animals benefited but the humans received little benefit or harm. Those animals that were most capable of taking advantage of the resources associated with human camps would have been the ‘tamer’ individuals: less aggressive, with shorter fight-or-flight distances. Later, these animals developed closer social or economic bonds with humans and led to a domestic relationship.[5][6]

The leap from a synanthropic population to a domestic one could only have taken place after the animals had progressed from anthropophily to habituation, to commensalism and partnership, at which point the establishment of a reciprocal relationship between animal and human would have laid the foundation for domestication, including captivity and human-controlled breeding. From this perspective, animal domestication is a coevolutionary process in which a population responds to selective pressure while adapting to a novel niche that includes another species with evolving behaviors.[6]

Commensal pathway animals include dogs, cats, fowl, and possibly pigs.

Dogs

The dog was the first domesticated animal, and was domesticated and widely established across Eurasia before the end of the Pleistocene, well before the cultivation of crops or the domestication of other animals.[7] The dog is often hypothesised to be a classic example of a domestic animal that likely traveled a commensal pathway into domestication. Archaeological evidence, such as the Bonn-Oberkassel dog dating to ~14,000BP,[8] supports the hypothesis that dog domestication preceded the emergence of agriculture [9][10] and began close to the Last Glacial Maximum when hunter-gatherers preyed on megafauna. The wolves more likely drawn to human camps were the less-aggressive, subdominant pack members with lowered flight response, higher stress thresholds, and less wary around humans, and therefore better candidates for domestication.[5] Proto-dogs might have taken advantage of carcasses left on site by early hunters, assisted in the capture of prey, or provided defense from large competing predators at kills.[10] However, the extent to which proto-domestic wolves could have become dependent on this way of life prior to domestication and without human provisioning is unclear and highly debated. In contrast, cats may have become fully dependent on a commensal lifestyle before being domesticated by preying on other commensal animals, such as rats and mice, without any human provisioning. Debate over the extent to which some wolves were commensal with humans prior to domestication stems from debate over the level of human intentionality in the domestication process, which remains untested.[11][12]

The earliest sign of domestication in dogs was the neotonization of skull morphology[13][14][5] and the shortening of snout length that results in tooth crowding, reduction in tooth size, and a reduction in the number of teeth,[15][5] which has been attributed to the strong selection for reduced aggression.[14][5] This process may have begun during the initial commensal stage of dog domestication, even before humans began to be active partners in the process.[5][6]

A mitochondrial, microsatellite, and Y-chromosome assessment of two wolf populations in North America combined with satellite telemetry data revealed significant genetic and morphological differences between one population that migrated with and preyed upon caribou and another territorial ecotype population that remained in a boreal coniferous forest. Although these two populations spend a period of the year in the same place, and though there was evidence of gene flow between them, the difference in prey–habitat specialization has been sufficient to maintain genetic and even coloration divergence.[16][6]

A different study has identified the remains of a population of extinct Pleistocene Beringian wolves with unique mitochondrial signatures. The skull shape, tooth wear, and isotopic signatures suggested these remains were derived from a population of specialist megafauna hunters and scavengers that became extinct while less specialized wolf ecotypes survived.[17][6] Analogous to the modern wolf ecotype that has evolved to track and prey upon caribou, a Pleistocene wolf population could have begun following mobile hunter-gatherers, thus slowly acquiring genetic and phenotypic differences that would have allowed them to more successfully adapt to the human habitat.[18][6]

Aspergillus and Staphylococcus

Numerous genera of bacteria and fungi live on and in the human body as part of its natural flora. The fungal genus Aspergillus is capable of living under considerable environmental stress, and thus is capable of colonising the upper gastrointestinal tract where relatively few examples of the body's gut flora can survive due to highly acidic or alkaline conditions produced by gastric acid and digestive juices. While Aspergillus normally produces no symptoms, in individuals who are immunocompromised or suffering from existing conditions such as tuberculosis, a condition called aspergillosis can occur, in which populations of Aspergillus grow out of control.

Staphylococcus aureus, a common bacterial species, is known best for its numerous pathogenic strains that can cause numerous illnesses and conditions. However, many strains of S. aureus are metabiotic commensals, and are present on roughly 20 to 30% of the human population as part of the skin flora.[19] S. aureus also benefits from the variable ambient conditions created by the body's mucous membranes, and as such can be found in the oral and nasal cavities, as well as inside the ear canal. Other Staphylococcus species including S. warneri, S. lugdunensis and S. epidermidis, will also engage in commensalism for similar purposes.

Arguments

Whether the relationship between humans and some types of gut flora is commensal or mutualistic is still unanswered.

Some biologists argue that any close interaction between two organisms is unlikely to be completely neutral for either party, and that relationships identified as commensal are likely mutualistic or parasitic in a subtle way that has not been detected. For example, epiphytes are "nutritional pirates" that may intercept substantial amounts of nutrients that would otherwise go to the host plant.[20] Large numbers of epiphytes can also cause tree limbs to break or shade the host plant and reduce its rate of photosynthesis. Similarly, phoretic mites may hinder their host by making flight more difficult, which may affect its aerial hunting ability or cause it to expend extra energy while carrying these passengers.

Types

Fly June 2008-2
Phoretic mites on a fly (Pseudolynchia canariensis)

Like all ecological interactions, commensalisms vary in strength and duration from intimate, long-lived symbioses to brief, weak interactions through intermediaries.

Phoresy

Phoresy is one animal attached to another exclusively for transport, mainly arthropods, examples of which are mites on insects (such as beetles, flies or bees), pseudoscorpions on mammals[21] or beetles, and millipedes on birds.[22] Phoresy can be either obligate or facultative (induced by environmental conditions).

Inquilinism

Tillandsia bourgaei
Inquilinism: Tillandsia bourgaei growing on an oak tree in Mexico

Inquilinism is the use of a second organism for permanent housing. Examples are epiphytic plants (such as many orchids) that grow on trees,[23] or birds that live in holes in trees.

Metabiosis

Metabiosis is a more indirect dependency, in which one organism creates or prepares a suitable environment for a second. Examples include maggots, which feast and develop on corpses, and hermit crabs, which use gastropod shells to protect their bodies.

See also

  • Mutualism – where both organisms experience mutual benefit in the relationship
  • Parasitism – where one organism benefits at the expense of another organism.
  • Parabiosis – where both organisms occupy the same dwelling, but do not interfere with each other
  • Symbiosis – long-term interactions between different biological species, which can be mutualistic, commensal or parasitic

References

  1. ^ Wilson, Edward O. (1975). "Ch.17-Social Symbiosis". Sociobiology: The New Synthesis. Harvard University Press. p. 354. ISBN 0-674-00089-7.
  2. ^ Mikula, Peter; Hadrava, Jiří; Albrecht, Tomáš; Tryjanowski, Piotr (19 March 2018). "Large-scale assessment of commensalistic–mutualistic associations between African birds and herbivorous mammals using internet photos". PeerJ. 6: e4520. doi:10.7717/peerj.4520. PMC 5863707. PMID 29576981.
  3. ^ Harper, Douglas. "commensalism". Online Etymology Dictionary.
  4. ^ van Beneden, Pierre-Joseph (1876). Animal parasites and messmates. London, Henry S. King. link.
  5. ^ a b c d e f Zeder MA (2012). "The domestication of animals". Journal of Anthropological Research. 68: 161–190. doi:10.3998/jar.0521004.0068.201.
  6. ^ a b c d e f Larson, G (2014). "The Evolution of Animal Domestication" (PDF). Annual Review of Ecology, Evolution, and Systematics. 45: 115–36. doi:10.1146/annurev-ecolsys-110512-135813.
  7. ^ Larson G (2012). "Rethinking dog domestication by integrating genetics, archeology, and biogeography" (PDF). Proceedings of the National Academy of Sciences of the United States of America. 109: 8878–83. Bibcode:2012PNAS..109.8878L. doi:10.1073/pnas.1203005109. PMC 3384140. PMID 22615366.
  8. ^ Janssens, L. (2018). "A new look at an old dog: Bonn-Oberkassel reconsidered". Journal of Archaeological Science. 92: 126–138. doi:10.1016/j.jas.2018.01.004.
  9. ^ Vila, C. (1997). "Multiple and ancient origins of the domestic dog". Science. 276 (5319): 1687–9. doi:10.1126/science.276.5319.1687. PMID 9180076.
  10. ^ a b Thalmann, O. (2013). "Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs". Science. 342: 871–874. Bibcode:2013Sci...342..871T. doi:10.1126/science.1243650. PMID 24233726.
  11. ^ Larson G, Fuller DQ (2014). "The evolution of animal domestication". Annual Review of Ecology, Evolution, and Systematics. 45: 217–295. doi:10.1146/annurev-ecolsys-110512-135813.
  12. ^ Hulme-Beaman A, Dobney K, Cucchi T, Searle JB (2016). "An ecological and evolutionary framework for commensalism in anthropogenic environments". Trends in Ecology and Evolution. 31: 633–645. doi:10.1016/j.tree.2016.05.001.CS1 maint: Multiple names: authors list (link)
  13. ^ Morey, Darcy F. 1992. Size, shape, and development in the evolution of the domestic dog. Journal of Archaeological Science 19:181–204
  14. ^ a b Lyudmila N. Trut (1999). "Early Canid Domestication: The Farm-Fox Experiment" (PDF). American Scientist. Sigma Xi, The Scientific Research Society. 87 (March–April): 160–169. Bibcode:1999AmSci..87.....T. doi:10.1511/1999.20.813. Retrieved June 25, 2011.
  15. ^ Turnbull, Priscilla F., and Charles A. Reed. 1974. The fauna from the terminal Pleistocene of Palegawra Cave. Fieldiana: Anthropology 63:81–146
  16. ^ Musiani M, Leonard JA, Cluff H, Gates CC, Mariani S, et al. (2007). "Differentiation of tundra/taiga and boreal coniferous forest wolves: genetics, coat colour and association with migratory caribou". Mol. Ecol. 16: 4149–70. doi:10.1111/j.1365-294x.2007.03458.x. PMID 17725575.
  17. ^ Leonard, J.A. (2007). "Megafaunal extinctions and the disappearance of a specialized wolf ecomorph" (PDF). Current Biology. 17 (13): 1146–50. doi:10.1016/j.cub.2007.05.072. PMID 17583509.
  18. ^ Wolpert, Stuart (November 14, 2013). "Dogs likely originated in Europe more than 18,000 years ago, UCLA biologists report". UCLA News Room. Retrieved December 10, 2014. Statement by Wayne, R.K.
  19. ^ Kluytmans J, van Belkum A, Verbrugh H (July 1997). "Nasal carriage of Staphylococcus aureus: epidemiology, underlying mechanisms, and associated risks". Clin. Microbiol. Rev. 10 (3): 505–20. PMC 172932. PMID 9227864.
  20. ^ Benzing, D.H. (1980) Biology of the Bromeliads. Eureka, California: Mad River Press.
  21. ^ Durden, Lance A. (June 1991). "Pseudoscorpions Associated With Mammals in Papua New Guinea". Biotropica. 23 (2): 204–6. doi:10.2307/2388309. JSTOR 2388309.
  22. ^ Tajovský, Karel; Mock, Andrej; Krumpál, Miroslav (2001). "Millipedes (Diplopoda) in birdsˈ nests". European Journal of Soil Biology. 37 (4): 321–3. doi:10.1016/S1164-5563(01)01108-6.
  23. ^ C. Michael Hogan. 2011. Commensalism. Topic Ed. M.Mcginley. Ed-in-chief C.J.Cleveland. Encyclopedia of Earth. National Council for Science and the Environment. Washington DC

External links

Bacterivore

Bacterivores are free-living, generally heterotrophic organisms, exclusively microscopic, which obtain energy and nutrients primarily or entirely from the consumption of bacteria. Many species of amoeba are bacterivores, as well as other types of protozoans. Commonly, all species of bacteria will be prey, but spores of some species, such as Clostridium perfringens, will never be prey, because of their cellular attributes.

Biological interaction

In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species (intraspecific interactions), or of different species (interspecific interactions). These effects may be short-term, like pollination and predation, or long-term; both often strongly influence the evolution of the species involved. A long-term interaction is called a symbiosis. Symbioses range from mutualism, beneficial to both partners, to competition, harmful to both partners. Interactions can be indirect, through intermediaries such as shared resources or common enemies.

Community (ecology)

In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area and in a particular time, also known as a biocoenosis. The term community has a variety of uses. In its simplest form it refers to groups of organisms in a specific place or time, for example, "the fish community of Lake Ontario before industrialization".

Community ecology or synecology is the study of the interactions between species in communities on many spatial and temporal scales, including the distribution, structure, abundance, demography, and interactions between coexisting populations. The primary focus of community ecology is on the interactions populations as determined by specific genotypic and phenotypic characteristics. Community ecology has its origin in European plant sociology. Modern community ecology examines patterns such as variation in species richness, equitability, productivity and food web structure (see community structure); it also examines processes such as predator–prey population dynamics, succession, and community assembly.

On a deeper level the meaning and value of the community concept in ecology is up for debate. Communities have traditionally been understood on a fine scale in terms of local processes constructing (or destructing) an assemblage of species, such as the way climate change is likely to affect the make-up of grass communities. Recently this local community focus has been criticised. Robert Ricklefs has argued that it is more useful to think of communities on a regional scale, drawing on evolutionary taxonomy and biogeography, where some species or clades evolve and others go extinct.

Consumer (food chain)

Consumers are organisms that eat organisms from a different population. These organisms are formally referred to as heterotrophs, which include animals, some bacteria and fungi. Such organisms may consume by various means, they are called primary consumers.

Demodex folliculorum

Demodex folliculorum is a microscopic mite that can only survive on the skin of humans. Most people have D. folliculorum on their skin. Usually, the mites do not cause any harm, and are therefore considered an example of commensalism rather than parasitism. If D. folliculorum does cause disease, this is known as demodicosis.

Dominance (ecology)

Ecological dominance is the degree to which a taxon is more numerous than its competitors in an ecological community, or makes up more of the biomass. Most ecological communities are defined by their dominant species.

In many examples of wet woodland in western Europe, the dominant tree is alder (Alnus glutinosa).

In temperate bogs, the dominant vegetation is usually species of Sphagnum moss.

Tidal swamps in the tropics are usually dominated by species of mangrove (Rhizophoraceae)

Some sea floor communities are dominated by brittle stars.

Exposed rocky shorelines are dominated by sessile organisms such as barnacles and limpets.

Ectosymbiosis

Ectosymbiosis is form of symbiotic behavior in which a parasite lives on the body surface of the host, including internal surfaces such as the lining of the digestive tube and the ducts of glands. The parasitic species is generally an immobile, or sessile, organism existing off of biotic substrate through mutualism, commensalism, or parasitism. Ectosymbiosis is found throughout a diverse array of environments and in many different species.

In some species the symbiotic environment provided by both the parasite and host are mutually beneficial. In recent research it has been found that these micro-flora will evolve and diversify rapidly in response to a change in the external environment, in order to stabilize and maintain a beneficial ectosymbiotic environment.

Feeding frenzy

In ecology, a feeding frenzy occurs when predators are overwhelmed by the amount of prey available. For example, a large school of fish can cause nearby sharks, such as the lemon shark, to enter into a feeding frenzy. This can cause the sharks to go wild, biting anything that moves, including each other or anything else within biting range. Another functional explanation for feeding frenzy is competition amongst predators. This term is most often used when referring to sharks or piranhas. It has also been used as a term within journalism.

Mesotrophic soil

Mesotrophic soils are soils with a moderate inherent fertility. An indicator of soil fertility is its base status, which is expressed as a ratio relating the major nutrient cations (calcium, magnesium, potassium and sodium) found there to the soil's clay percentage. This is commonly expressed in hundredths of a mole of cations per kilogram of clay, i.e. cmol (+) kg−1 clay.

Non-trophic networks

Any action or influence that species have on each other is considered a biological interaction. These interactions between species can be considered in several ways. One such way is to depict interactions in the form of a network, which identifies the members and the patterns that connect them. Species interactions are considered primarily in terms of trophic interactions, which depict which species feed on others.

Currently, ecological networks that integrate non-trophic interactions are being built. The type of interactions they can contain can be classified into six categories: mutualism, commensalism, neutralism, amensalism, antagonism, and competition.

Observing and estimating the fitness costs and benefits of species interactions can be very problematic. The way interactions are interpreted can profoundly affect the ensuing conclusions.

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.

Pleocola limnoriae

Pleocola limnoriae is a species of tardigrades. It is the only species of the genus Pleocola, which belongs to the family Styraconyxidae. The species and genus were named by Alexandre Cantacuzène in 1951. It has been found in the North Atlantic Ocean, type locality Roscoff, Brittany, France. It was discovered living in commensalism on the isopod Limnoria lignorum.

Portuguese man o' war

The Atlantic Portuguese man o' war (Physalia physalis), also known as the man-of-war, is a marine hydrozoan of the family Physaliidae found in the Atlantic, Indian and Pacific Oceans. Its long tentacles deliver a painful sting, which is venomous and powerful enough to kill fish or, rarely, humans. Despite its appearance, the Portuguese man o' war is not a true jellyfish but a siphonophore, which is not actually a single multicellular organism (true jellyfish are single organisms), but a colonial organism made up of specialized individual animals (of the same species) called zooids or polyps. These polyps are attached to one another and physiologically integrated, to the extent that they cannot survive independently, creating a symbiotic relationship, requiring each polyp to work together and function like an individual animal.

The Indo-Pacific Portuguese man-of-war (P. utriculus), or blue bottle, is a related species with very similar appearance found throughout the Indian and Pacific Oceans.

Pseudoscorpion

A pseudoscorpion, also known as a false scorpion or book scorpion, is an arachnid belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida.

Pseudoscorpions are generally beneficial to humans since they prey on clothes moth larvae, carpet beetle larvae, booklice, ants, mites, and small flies. They are tiny and inoffensive, and are rarely seen due to their small size, despite being common in many environments. Pseudoscorpions often carry out phoresy, a form of commensalism in which one organism uses another for the purpose of transport.

Recruitment (biology)

In biology, especially marine biology, recruitment occurs when a juvenile organism joins a population, whether by birth or immigration, usually at a stage whereby the organisms are settled and able to be detected by an observer.There are two types of recruitment: closed and open.In the study of fisheries, recruitment is "the number of fish surviving to enter the fishery or to some life history stage such as settlement or maturity".

Sarracenia purpurea

Sarracenia purpurea, commonly known as the purple pitcher plant, northern pitcher plant, turtle socks, or side-saddle flower, is a carnivorous plant in the family Sarraceniaceae.

Sea cucumber

Sea cucumbers are echinoderms from the class Holothuroidea. They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothurian () species worldwide is about 1,717 with the greatest number being in the Asia Pacific region. Many of these are gathered for human consumption and some species are cultivated in aquaculture systems. The harvested product is variously referred to as trepang, namako, bêche-de-mer or balate. Sea cucumbers serve a useful role in the marine ecosystem as they help recycle nutrients, breaking down detritus and other organic matter after which bacteria can continue the degradation process.Like all echinoderms, sea cucumbers have an endoskeleton just below the skin, calcified structures that are usually reduced to isolated microscopic ossicles (or sclerietes) joined by connective tissue. In some species these can sometimes be enlarged to flattened plates, forming an armour. In pelagic species such as Pelagothuria natatrix (Order Elasipodida, family Pelagothuriidae), the skeleton is absent and there is no calcareous ring.The sea cucumbers are named after their resemblance to the fruit of the cucumber plant.

Symbiosis

Symbiosis (from Greek συμβίωσις "living together", from σύν "together" and βίωσις "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasitic. The organisms, each termed a symbiont, may be of the same or of different species. In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term was subject to a century-long debate about whether it should specifically denote mutualism, as in lichens; biologists have now abandoned that restriction.

Symbiosis can be obligatory, which means that one or both of the symbionts entirely depend on each other for survival, or facultative (optional) when they can generally live independently.

Symbiosis is also classified by physical attachment; symbiosis in which the organisms have bodily union is called conjunctive symbiosis, and symbiosis in which they are not in union is called disjunctive symbiosis.

When one organism lives on the surface of another, such as head lice on humans, it is called ectosymbiosis; when one partner lives inside the tissues of another, such as Symbiodinium within coral, it is termed endosymbiosis.

Velleius dilatatus

The rove beetle Velleius dilatatus lives together with the European hornet Vespa crabro crabro.

Velleius dilatatus is up to 26 mm long. It feeds on detritus produced by the hornets (an example of commensalism). The beetles cannot survive without the hornets themselves, even if there is enough detritus present.

It finds hornet nests by its very good sense of smell. Most nests contain an average of ten V. dilatatus.

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