It has been suggested that Protocooperation be merged into this article. (Discuss) Proposed since November 2024. |
Mutualism describes the ecological interaction between two or more species where each species has a net benefit. [1] Mutualism is a common type of ecological interaction. Prominent examples are:
Mutualism can be contrasted with interspecific competition, in which each species experiences reduced fitness, and exploitation, and with parasitism, in which one species benefits at the expense of the other. [2] However, mutualism may evolve from interactions that began with imbalanced benefits, such as parasitism. [3]
The term mutualism was introduced by Pierre-Joseph van Beneden in his 1876 book Animal Parasites and Messmates to mean "mutual aid among species". [4] [5]
Mutualism is often conflated with two other types of ecological phenomena: cooperation and symbiosis. Cooperation most commonly refers to increases in fitness through within-species (intraspecific) interactions, although it has been used (especially in the past) to refer to mutualistic interactions, and it is sometimes used to refer to mutualistic interactions that are not obligate. [1] Symbiosis involves two species living in close physical contact over a long period of their existence and may be mutualistic, parasitic, or commensal, so symbiotic relationships are not always mutualistic, and mutualistic interactions are not always symbiotic. Despite a different definition between mutualism and symbiosis, they have been largely used interchangeably in the past, and confusion on their use has persisted. [6]
Mutualism plays a key part in ecology and evolution. For example, mutualistic interactions are vital for terrestrial ecosystem function as:
A prominent example of pollination mutualism is with bees and flowering plants. Bees use these plants as their food source with pollen and nectar. In turn, they transfer pollen to other nearby flowers, inadvertently allowing for cross-pollination. Cross-pollination has become essential in plant reproduction and fruit/seed production. The bees get their nutrients from the plants, and allow for successful fertilization of plants, demonstrating a mutualistic relationship between two seemingly-unlike species.
Mutualism has also been linked to major evolutionary events, such as the evolution of the eukaryotic cell (symbiogenesis) and the colonization of land by plants in association with mycorrhizal fungi.
Mutualistic relationships can be thought of as a form of "biological barter" [10] in mycorrhizal associations between plant roots and fungi, with the plant providing carbohydrates to the fungus in return for primarily phosphate but also nitrogenous compounds. Other examples include rhizobia bacteria that fix nitrogen for leguminous plants (family Fabaceae) in return for energy-containing carbohydrates. [11] Metabolite exchange between multiple mutualistic species of bacteria has also been observed in a process known as cross-feeding. [12] [13]
Service-resource relationships are common. Three important types are pollination, cleaning symbiosis, and zoochory.
In pollination, a plant trades food resources in the form of nectar or pollen for the service of pollen dispersal. However, daciniphilous Bulbophyllum orchid species trade sex pheromone precursor or booster components via floral synomones/attractants in a true mutualistic interactions with males of Dacini fruit flies (Diptera: Tephritidae: Dacinae). [14] [15]
Phagophiles feed (resource) on ectoparasites, thereby providing anti-pest service, as in cleaning symbiosis. Elacatinus and Gobiosoma , genera of gobies, feed on ectoparasites of their clients while cleaning them. [16]
Zoochory is the dispersal of the seeds of plants by animals. This is similar to pollination in that the plant produces food resources (for example, fleshy fruit, overabundance of seeds) for animals that disperse the seeds (service). Plants may advertise these resources using colour [17] and a variety of other fruit characteristics, e.g., scent. Fruit of the aardvark cucumber (Cucumis humifructus) is buried so deeply that the plant is solely reliant upon the aardvark's keen sense of smell to detect its ripened fruit, extract, consume and then scatter its seeds; [18] C. humifructus's geographical range is thus restricted to that of the aardvark's.
Another type is ant protection of aphids, where the aphids trade sugar-rich honeydew (a by-product of their mode of feeding on plant sap) in return for defense against predators such as ladybugs.[ citation needed ]
Strict service-service interactions are very rare, for reasons that are far from clear. [10] One example is the relationship between sea anemones and anemone fish in the family Pomacentridae: the anemones provide the fish with protection from predators (which cannot tolerate the stings of the anemone's tentacles) and the fish defend the anemones against butterflyfish (family Chaetodontidae), which eat anemones. However, in common with many mutualisms, there is more than one aspect to it: in the anemonefish-anemone mutualism, waste ammonia from the fish feeds the symbiotic algae that are found in the anemone's tentacles. [19] [20] Therefore, what appears to be a service-service mutualism in fact has a service-resource component. A second example is that of the relationship between some ants in the genus Pseudomyrmex and trees in the genus Acacia , such as the whistling thorn and bullhorn acacia. The ants nest inside the plant's thorns. In exchange for shelter, the ants protect acacias from attack by herbivores (which they frequently eat when those are small enough, introducing a resource component to this service-service relationship) and competition from other plants by trimming back vegetation that would shade the acacia. In addition, another service-resource component is present, as the ants regularly feed on lipid-rich food-bodies called Beltian bodies that are on the Acacia plant. [21]
In the neotropics, the ant Myrmelachista schumanni makes its nest in special cavities in Duroia hirsute . Plants in the vicinity that belong to other species are killed with formic acid. This selective gardening can be so aggressive that small areas of the rainforest are dominated by Duroia hirsute. These peculiar patches are known by local people as "devil's gardens". [22]
In some of these relationships, the cost of the ant's protection can be quite expensive. Cordia sp. trees in the Amazonian rainforest have a kind of partnership with Allomerus sp. ants, which make their nests in modified leaves. To increase the amount of living space available, the ants will destroy the tree's flower buds. The flowers die and leaves develop instead, providing the ants with more dwellings. Another type of Allomerus sp. ant lives with the Hirtella sp. tree in the same forests, but in this relationship, the tree has turned the tables on the ants. When the tree is ready to produce flowers, the ant abodes on certain branches begin to wither and shrink, forcing the occupants to flee, leaving the tree's flowers to develop free from ant attack. [22]
The term "species group" can be used to describe the manner in which individual organisms group together. In this non-taxonomic context one can refer to "same-species groups" and "mixed-species groups." While same-species groups are the norm, examples of mixed-species groups abound. For example, zebra ( Equus burchelli ) and wildebeest ( Connochaetes taurinus ) can remain in association during periods of long distance migration across the Serengeti as a strategy for thwarting predators. Cercopithecus mitis and Cercopithecus ascanius , species of monkey in the Kakamega Forest of Kenya, can stay in close proximity and travel along exactly the same routes through the forest for periods of up to 12 hours. These mixed-species groups cannot be explained by the coincidence of sharing the same habitat. Rather, they are created by the active behavioural choice of at least one of the species in question. [23]
[Mutualistic symbiosis can sometimes evolve from parasitism or commensalism. Symbiogenesis, a leading theory on the evolution of Eukaryotes states the origin of the mitochondria and cell nucleus emerged from a parasitic relationship of ancient Archaea and Bacteria. Fungi's relationship to plants in the form of mycelium evolved from parasitism and commensalism. Under certain conditions species of fungi previously in a state of mutualism can turn parasitic on weak or dying plants. [24] Likewise the symbiotic relationship of clown fish and sea anemones emerged from a commensalist relationship. [25] [26] [27] Once a mutualistic relationship emerges both symbionts are pushed towards co-evolution with each other. [28] [29]
Mathematical treatments of mutualisms, like the study of mutualisms in general, have lagged behind those for predation, or predator-prey, consumer-resource, interactions. In models of mutualisms, the terms "type I" and "type II" functional responses refer to the linear and saturating relationships, respectively, between the benefit provided to an individual of species 1 (dependent variable) and the density of species 2 (independent variable).[ citation needed ]
One of the simplest frameworks for modeling species interactions is the Lotka–Volterra equations. [30] In this model, the changes in population densities of the two mutualists are quantified as:
where
Mutualism is in essence the logistic growth equation modified for mutualistic interaction. The mutualistic interaction term represents the increase in population growth of one species as a result of the presence of greater numbers of another species. As the mutualistic interactive term β is always positive, this simple model may lead to unrealistic unbounded growth. [31] So it may be more realistic to include a further term in the formula, representing a saturation mechanism, to avoid this occurring.
In 1989, David Hamilton Wright modified the above Lotka–Volterra equations by adding a new term, βM/K, to represent a mutualistic relationship. [32] Wright also considered the concept of saturation, which means that with higher densities, there is a decrease in the benefits of further increases of the mutualist population. Without saturation, depending on the size of parameter α, species densities would increase indefinitely. Because that is not possible due to environmental constraints and carrying capacity, a model that includes saturation would be more accurate. Wright's mathematical theory is based on the premise of a simple two-species mutualism model in which the benefits of mutualism become saturated due to limits posed by handling time. Wright defines handling time as the time needed to process a food item, from the initial interaction to the start of a search for new food items and assumes that processing of food and searching for food are mutually exclusive. Mutualists that display foraging behavior are exposed to the restrictions on handling time. Mutualism can be associated with symbiosis.[ citation needed ]
In 1959, C. S. Holling performed his classic disc experiment that assumed that
where
The equation that incorporates Type II functional response and mutualism is:
where
or, equivalently,
where
This model is most effectively applied to free-living species that encounter a number of individuals of the mutualist part in the course of their existences. Wright notes that models of biological mutualism tend to be similar qualitatively, in that the featured isoclines generally have a positive decreasing slope, and by and large similar isocline diagrams. Mutualistic interactions are best visualized as positively sloped isoclines, which can be explained by the fact that the saturation of benefits accorded to mutualism or restrictions posed by outside factors contribute to a decreasing slope.
The type II functional response is visualized as the graph of vs.M.
Mutualistic networks made up out of the interaction between plants and pollinators were found to have a similar structure in very different ecosystems on different continents, consisting of entirely different species. [33] The structure of these mutualistic networks may have large consequences for the way in which pollinator communities respond to increasingly harsh conditions and on the community carrying capacity. [34]
Mathematical models that examine the consequences of this network structure for the stability of pollinator communities suggest that the specific way in which plant-pollinator networks are organized minimizes competition between pollinators, [35] reduce the spread of indirect effects and thus enhance ecosystem stability [36] and may even lead to strong indirect facilitation between pollinators when conditions are harsh. [37] This means that pollinator species together can survive under harsh conditions. But it also means that pollinator species collapse simultaneously when conditions pass a critical point. [38] This simultaneous collapse occurs, because pollinator species depend on each other when surviving under difficult conditions. [37]
Such a community-wide collapse, involving many pollinator species, can occur suddenly when increasingly harsh conditions pass a critical point and recovery from such a collapse might not be easy. The improvement in conditions needed for pollinators to recover could be substantially larger than the improvement needed to return to conditions at which the pollinator community collapsed. [37]
Humans are involved in mutualisms with other species: their gut flora is essential for efficient digestion. [39] Infestations of head lice might have been beneficial for humans by fostering an immune response that helps to reduce the threat of body louse borne lethal diseases. [40]
Some relationships between humans and domesticated animals and plants are to different degrees mutualistic. For example, agricultural varieties of maize provide food for humans and are unable to reproduce without human intervention because the leafy sheath does not fall open, and the seedhead (the "corn on the cob") does not shatter to scatter the seeds naturally.[ citation needed ]
In traditional agriculture, some plants have mutualist as companion plants, providing each other with shelter, soil fertility and/or natural pest control. For example, beans may grow up cornstalks as a trellis, while fixing nitrogen in the soil for the corn, a phenomenon that is used in Three Sisters farming. [41]
One researcher has proposed that the key advantage Homo sapiens had over Neanderthals in competing over similar habitats was the former's mutualism with dogs. [42]
The microbiota in the human intestine coevolved with the human species, and this relationship is considered to be a mutualism that is beneficial both to the human host and the bacteria in the gut population. [43] The mucous layer of the intestine contains commensal bacteria that produce bacteriocins, modify the pH of the intestinal contents, and compete for nutrition to inhibit colonization by pathogens. [44] The gut microbiota, containing trillions of microorganisms, possesses the metabolic capacity to produce and regulate multiple compounds that reach the circulation and act to influence the function of distal organs and systems. [45] Breakdown of the protective mucosal barrier of the gut can contribute to the development of colon cancer. [44]
Every generation of every organism needs nutrients – and similar nutrients – more than they need particular defensive characteristics, as the fitness benefit of these vary heavily especially by environment. This may be the reason that hosts are more likely to evolve to become dependent on vertically transmitted bacterial mutualists which provide nutrients than those providing defensive benefits. This pattern is generalized beyond bacteria by Yamada et al. 2015's demonstration that undernourished Drosophila are heavily dependent on their fungal symbiont Issatchenkia orientalis for amino acids. [46]
Mutualisms are not static, and can be lost by evolution. [47] Sachs and Simms (2006) suggest that this can occur via four main pathways:
There are many examples of mutualism breakdown. For example, plant lineages inhabiting nutrient-rich environments have evolutionarily abandoned mycorrhizal mutualisms many times independently. [50] Evolutionarily, headlice may have been mutualistic as they allow for early immunity to various body-louse borne disease; however, as these diseases became eradicated, the relationship has become less mutualistic and more parasitic. [48]
Measuring the exact fitness benefit to the individuals in a mutualistic relationship is not always straightforward, particularly when the individuals can receive benefits from a variety of species, for example most plant-pollinator mutualisms. It is therefore common to categorise mutualisms according to the closeness of the association, using terms such as obligate and facultative. Defining "closeness", however, is also problematic. It can refer to mutual dependency (the species cannot live without one another) or the biological intimacy of the relationship in relation to physical closeness (e.g., one species living within the tissues of the other 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".
A mycorrhiza is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play important roles in plant nutrition, soil biology, and soil chemistry.
Ectosymbiosis is a form of symbiotic behavior in which an organism lives on the body surface of another organism, 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 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 biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.
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.
Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In general, they are gram negative, motile, non-sporulating rods.
Microbial ecology is the ecology of microorganisms: their relationship with one another and with their environment. It concerns the three major domains of life—Eukaryota, Archaea, and Bacteria—as well as viruses. This relationship is often mediated by secondary metabolites produced by microorganisms. These secondary metabolites are known as specialized metabolites and are mostly volatile or non volatile compounds. These metabolites include terpenoids, sulfur compounds, indole compound and many more.
The Prodoxidae are a family of moths, generally small in size and nondescript in appearance. They include species of moderate pest status, such as the currant shoot borer, and others of considerable ecological and evolutionary interest, such as various species of "yucca moths".
Myrmecophytes are plants that live in a mutualistic association with a colony of ants. There are over 100 different genera of myrmecophytes. These plants possess structural adaptations in the form of domatia where ants can shelter, and food bodies and extrafloral nectaries that provide ants with food. In exchange for these resources, ants aid the myrmecophyte in pollination, seed dispersal, gathering of essential nutrients, and defense. Domatia adapted specifically to ants may be called myrmecodomatia.
Myrmecochory ( ; from Ancient Greek: μύρμηξ, romanized: mýrmēks and χορεία khoreíā is seed dispersal by ants, an ecologically significant ant–plant interaction with worldwide distribution. Most myrmecochorous plants produce seeds with elaiosomes, a term encompassing various external appendages or "food bodies" rich in lipids, amino acids, or other nutrients that are attractive to ants. The seed with its attached elaiosome is collectively known as a diaspore. Seed dispersal by ants is typically accomplished when foraging workers carry diaspores back to the ant colony, after which the elaiosome is removed or fed directly to ant larvae. Once the elaiosome is consumed, the seed is usually discarded in an underground midden or ejected from the nest. Although diaspores are seldom distributed far from the parent plant, myrmecochores also benefit from this predominantly mutualistic interaction through dispersal to favourable locations for germination, as well as escape from seed predation.
Ant–fungus mutualism is a symbiosis seen between certain ant and fungal species, in which ants actively cultivate fungus much like humans farm crops as a food source. There is only evidence of two instances in which this form of agriculture evolved in ants resulting in a dependence on fungi for food. These instances were the attine ants and some ants that are part of the Megalomyrmex genus. In some species, the ants and fungi are dependent on each other for survival. This type of codependency is prevalent among herbivores who rely on plant material for nutrition. The fungus’ ability to convert the plant material into a food source accessible to their host makes them the ideal partner. The leafcutter ant is a well-known example of this symbiosis. Leafcutter ants species can be found in southern South America up to the United States. However, ants are not the only ground-dwelling arthropods which have developed symbioses with fungi. A similar mutualism with fungi is also noted in termites within the subfamily Macrotermitinae which are widely distributed throughout the Old World tropics with the highest diversity in Africa.
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.
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 (semiochemical) in the environment by insects. Semiochemicals are secreted by the organisms in the environment and they are detected by other organism such as insects. Semiochemicals 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 pheromone, synomones, allomone and kairomone. Pheromones are semiochemicals that facilitates interaction between organisms of same species. Synomones benefit both the producer and receiver, allomone is advantageous to only the producer whiles kairomones is beneficial to the receiver. Insect interact with other species within their community and these interaction include mutualism, commensalism, ammensalism, parasitism and neutralisms.
An ecological network is a representation of the biotic interactions in an ecosystem, in which species (nodes) are connected by pairwise interactions (links). These interactions can be trophic or symbiotic. Ecological networks are used to describe and compare the structures of real ecosystems, while network models are used to investigate the effects of network structure on properties such as ecosystem stability.
Upiga is a monotypic moth genus described by Hahn William Capps in 1964. The genus is placed in the family Crambidae, but has also been placed in Pyralidae. It contains only one species, Upiga virescens, the senita moth, described by George Duryea Hulst in 1900 and found in the Sonoran Desert of North America.
A pollination network is a bipartite mutualistic network in which plants and pollinators are the nodes, and the pollination interactions form the links between these nodes. The pollination network is bipartite as interactions only exist between two distinct, non-overlapping sets of species, but not within the set: a pollinator can never be pollinated, unlike in a predator-prey network where a predator can be depredated. A pollination network is two-modal, i.e., it includes only links connecting plant and animal communities.
Jordi Bascompte is a professor of ecology at the University of Zurich and the director of its specialized master's program on quantitative environmental sciences. He is best known for having brought the interactions of mutual benefit between plants and animals into community ecology, at the time largely dominated by predation and competition. His application of network theory to the study of mutualism has identified general laws that determine the way in which species interactions shape biodiversity.
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.