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Ectosymbiosis is a form of symbiotic behavior in which an organism lives on the body surface of another organism (the host), including internal surfaces such as the lining of the digestive tube and the ducts of glands. The ectosymbiotic species, or ectosymbiont, is generally an immobile (or sessile) organism existing off of biotic substrate through mutualism, commensalism, or parasitism. [1] [2] 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. [3]
Ectosymbiosis has evolved independently many times to fill a wide variety of ecological niches, both temperate and extreme. [4] Such temperate regions include the seas off the coast of Singapore while the extreme regions reach to the depths of Antarctica and hydrothermal vents. [1] [2] [5] It likely evolved as a niche specialization, which allowed for greater diversity in ectosymbiotic behavior among species. Additionally, in the case of mutualism, the evolution improved the fitness of both species involved, propagating the success of ectosymbiosis. Ectosymbiosis has independently evolved through convergent evolution in all domains of life. [4] [6]
Ectosymbiosis allows niches to form that would otherwise be unable to exist without the support of their host. Inherently, this added niche opens up a new branch off of the evolutionary tree. The evolutionary success of ectosymbiosis is based on the benefits experienced by the ectosymbiont and the host. Due to the dependence of the parasite on the host and the associated benefits and cost to both the parasite and host, the two will continue to coevolve as explained by the Red Queen hypothesis. [7] The Red Queen hypothesis states that a host will continually evolve defenses against a parasitic attack, and the parasite species will also adapt to these changes in the host defense, the result being competitive coevolution between the two species. [7]
Ectosymbiosis adds to the biodiversity of the environment, whether on land, in freshwater, in deserts, or in deep sea vents. [8] Specifically, ectosymbiosis provides a new niche or environment from which many new species can differentiate and flourish.
This niche specialization between species also leads to stabilization of symbiotic relationships between sessile and motile organisms. The ectosymbiont can increase the fitness of their host by assisting with metabolism, nitrogen fixation, or cleaning the host organism. [3] [9] [10] The diversity of advantages has yet to be fully explored, but by virtue of persisting throughout all of recent evolution, they likely confer an adaptive advantage to many of the species that exist solely due to ectosymbiosis.
Although ectosymbiosis is typically an evolutionary stable behavior, the different host and parasite dynamics independently vary in their stability.
Commensalism is a form of symbiosis where one species is benefiting from the interactions between species and the other is neither helped nor harmed from the interaction. Ectosymbiotic commensalistic behavior is found frequently in organisms that attach themselves to larger species in order to move long distances or scavenge food easily; this is documented in remoras which attach to sharks to scavenge and travel. [11] An additional ectosymbiotic example of commensalism is the relationship between small sessile organisms and echinoids in the Southern ocean, where the echinoids provide substrate for the small organisms to grow and the echinoids remain unaffected. [8]
Mutualism is a form of ectosymbiosis where both the host and parasitic species benefit from the interaction. There are many examples of mutualistic ectosymbiosis that occur in nature. One such relationship is between Branchiobdellida and crayfish in which the Branchiobdellida acts as a bacterial gut cleaner for the crayfish species. [10] Another example is the iron-oxide associated chemoautotrophic bacteria found crusted to the gills of Rimicaris exoculata shrimp that provide the shrimp with vital organic material for their survival while simultaneously supporting the bacteria with different organic material that the bacterial cannot produce itself. [5] Groups of organisms – greater than a single pair of a host and parasite – can also form mutualistic ectosymbiotic interactions. Bark beetles can work in a dynamic mutualistic fashion with fungi and mites attached to their exoskeletons, both of which feed off of trees to provide vital energy to the beetles while the beetles provide necessary organic material to the fungi and mites to survive. [12] [13] In this case, the relationship between the fungi and mites is functional because while both do the same job, they are optimally functional at different temperatures. [12] [13]
Mutualistic interactions can be evolutionarily unstable because of the constant battle to maximize one's self-benefits. [14] This is due to the limited benefits offered to both the parasite and the host, with the possible outcome for at least one of the species to die out if the other species begins to take advantage of the other. [14] In the case that the mutualistic behavior persists for enough generations, the dynamic can evolve into parasitism, which is a more stable dynamic due to the increased benefit to the parasite that propagates the behavior. [14] In this case the parasite takes advantage of the previously mutualistic host and parasite dynamic, gaining greater benefits for itself. [14]
Parasitism is a form of symbiosis in which one species benefits from the interactions between species while the other organism is actively harmed. This is the most common form of ectosymbiotic interactions. One of the many examples of ectosymbiotic parasites includes head lice in humans, which feed on blood by attaching to a human's scalp. Additionally, mature Branchiobdellida bacteria act as a nutrient thief in the gut of crayfish species to exist. In these cases, the head lice and the Branchiobdellida are both parasites interacting with host species. [10]
Symbiosis is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can be either in a mutualistic, a commensalistic, or a parasitic relationship. In 1879, Heinrich Anton de Bary defined symbiosis as "the living together of unlike organisms".
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery; animals such as hookworms, lice, mosquitoes, and vampire bats; fungi such as honey fungus and the agents of ringworm; and plants such as mistletoe, dodder, and the broomrapes.
Mutualism describes the ecological interaction between two or more species where each species has a net benefit. Mutualism is a common type of ecological interaction. Prominent examples are:
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. 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 is harmed and the other benefits.
In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist guest (symbiont). The guest is typically provided with nourishment and shelter. Examples include animals playing host to parasitic worms, cells harbouring pathogenic (disease-causing) viruses, or a bean plant hosting mutualistic (helpful) nitrogen-fixing bacteria. More specifically in botany, a host plant supplies food resources to micropredators, which have an evolutionarily stable relationship with their hosts similar to ectoparasitism. The host range is the collection of hosts that an organism can use as a partner.
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, or of different species. These effects may be short-term, or long-term, both often strongly influence the adaptation and evolution of the species involved. Biological interactions range from mutualism, beneficial to both partners, to competition, harmful to both partners. Interactions can be direct when physical contact is established or indirect, through intermediaries such as shared resources, territories, ecological services, metabolic waste, toxins or growth inhibitors. This type of relationship can be shown by net effect based on individual effects on both organisms arising out of relationship.
The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microorganisms that feed on sloughed-off plant cells, termed rhizodeposition, and the proteins and sugars released by roots, termed root exudates. This symbiosis leads to more complex interactions, influencing plant growth and competition for resources. Much of the nutrient cycling and disease suppression by antibiotics required by plants occurs immediately adjacent to roots due to root exudates and metabolic products of symbiotic and pathogenic communities of microorganisms. The rhizosphere also provides space to produce allelochemicals to control neighbours and relatives.
Symbiotic bacteria are bacteria living in symbiosis with another organism or each other. For example, rhizobia living in root nodules of legumes provide nitrogen fixing activity for these plants.
Cheating is a term used in behavioral ecology and ethology to describe behavior whereby organisms receive a benefit at the cost of other organisms. Cheating is common in many mutualistic and altruistic relationships. A cheater is an individual who does not cooperate but can potentially gain the benefit from others cooperating. Cheaters are also those who selfishly use common resources to maximize their individual fitness at the expense of a group. Natural selection favors cheating, but there are mechanisms to regulate it. The stress gradient hypothesis states that facilitation, cooperation or mutualism should be more common in stressful environments, while cheating, competition or parasitism are common in benign environments.
Myrmecophily consists of positive, mutualistic, interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. It may also include commensal or even parasitic interactions.
In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community, or life assemblage. 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".
Insect ecology is the interaction of insects, individually or as a community, with the surrounding environment or ecosystem. This interaction is mostly mediated by the secretion and detection of chemicals (semiochemicals) in the environment by insects. Semiochemicals are secreted by the organisms in the environment and they are detected by other organism such as insects. Semiochemical used by organisms, including (insects) to interact with other organism either of the same species or different species can generally grouped into four. These are pheromones, synomones, allomones and kairomones. Pheromones are semiochemicals that facilitates interaction between organisms of same species. Synomones benefit both the producer and receiver, allomene is advantageous to only the producer whiles kairomones is beneficial to the receiver.
Cleaning symbiosis is a mutually beneficial association between individuals of two species, where one removes and eats parasites and other materials from the surface of the other. Cleaning symbiosis is well-known among marine fish, where some small species of cleaner fish, notably wrasses but also species in other genera, are specialised to feed almost exclusively by cleaning larger fish and other marine animals. Other cleaning symbioses exist between birds and mammals, and in other groups.
Evolving digital ecological networks are webs of interacting, self-replicating, and evolving computer programs that experience the same major ecological interactions as biological organisms. Despite being computational, these programs evolve quickly in an open-ended way, and starting from only one or two ancestral organisms, the formation of ecological networks can be observed in real-time by tracking interactions between the constantly evolving organism phenotypes. These phenotypes may be defined by combinations of logical computations that digital organisms perform and by expressed behaviors that have evolved. The types and outcomes of interactions between phenotypes are determined by task overlap for logic-defined phenotypes and by responses to encounters in the case of behavioral phenotypes. Biologists use these evolving networks to study active and fundamental topics within evolutionary ecology.
A lichenicolous fungus is a member of a specialised group of fungi that live exclusively on lichens as their host organisms. These fungi, comprising over 2,000 known species across 280 genera, exhibit a wide range of ecological strategies, including parasitism, commensalism, and mutualism. They can be found in diverse environments worldwide, from tropical to polar regions, and play important roles in lichen ecology and biodiversity. Lichenicolous fungi are classified into several taxonomic groups, with the majority belonging to the Ascomycota and a smaller portion to the Basidiomycota. Their interactions with host lichens range from mild parasitism to severe pathogenicity, sometimes causing significant damage to lichen communities.
Microbial symbiosis in marine animals was not discovered until 1981. In the time following, symbiotic relationships between marine invertebrates and chemoautotrophic bacteria have been found in a variety of ecosystems, ranging from shallow coastal waters to deep-sea hydrothermal vents. Symbiosis is a way for marine organisms to find creative ways to survive in a very dynamic environment. They are different in relation to how dependent the organisms are on each other or how they are associated. It is also considered a selective force behind evolution in some scientific aspects. The symbiotic relationships of organisms has the ability to change behavior, morphology and metabolic pathways. With increased recognition and research, new terminology also arises, such as holobiont, which the relationship between a host and its symbionts as one grouping. Many scientists will look at the hologenome, which is the combined genetic information of the host and its symbionts. These terms are more commonly used to describe microbial symbionts.
The mycobiome, mycobiota, or fungal microbiome, is the fungal community in and on an organism.
The Red King hypothesis contrasts with the Red Queen hypothesis, where mutualistic and cooperative interactions favor the fitness of a set of individuals through slow evolution, as opposed to having competitive interactions or having an "arms race". The hypothesis posits that individuals from different communities can establish positive interactions for long periods of time when there is a great benefit for both parties, also through mutual help, individuals from different species (communities) can share different tasks to build a niche, which avoid spending energy in competing and increasing their resilience over environmental stress.
Plant-animal interactions are important pathways for the transfer of energy within ecosystems, where both advantageous and unfavorable interactions support ecosystem health. Plant-animal interactions can take on important ecological functions and manifest in a variety of combinations of favorable and unfavorable associations, for example predation, frugivory and herbivory, parasitism, and mutualism. Without mutualistic relationships, some plants may not be able to complete their life cycles, and the animals may starve due to resource deficiency.
Obligate mutualism is a special case of mutualism where an ecological interaction between species mutually benefits each other, and one or all species are unable to survive without the other. In some obligate relationships, only one species is dependent on the relationship. For example, a parasite may require a host in order to reproduce and survive, while the host does not depend at all on the parasite. Fig and fig wasps are an example of a co-obligate relationship, where both species are totally dependent on the relationship. The fig plant is entirely dependent on the fig wasp for pollination, and the fig wasp requires the fig plant for reproductive purposes. Many insect-fungi relationships are also co-obligate: the insect disperses, and in some cases protects, the fungi while the fungi provide nutrients for the insects. This interaction allows insects and fungi to, as a group, inhabit previously inhospitable or unreachable environments. Though obligate relationships need not be limited to two species, they are often discussed as such, with the relationship being made up of a host and a symbiont, though the terms are often attributed arbitrarily.
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