ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...

, a biological interaction is the effect that a pair of

organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...

s living together in a

community A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, t ...

have on each other. They can be either of the same

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

(intraspecific interactions), or of different species (

interspecific interaction Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...

s). These effects may be short-term, like

pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...

and

predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...

, or long-term; both often strongly influence the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

of the species involved. A long-term interaction is called a

symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...

. Symbioses range from mutualism, beneficial to both partners, to

competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, ind ...

, harmful to both partners. Interactions can be indirect, through intermediaries such as shared resources or common enemies. This type of relationship can be shown by net effect based on individual effects on both organisms arising out of relationship. Several recent studies have suggested non-trophic species interactions such as habitat modification and mutualisms can be important determinants of food web structures. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups.

History

Although biological interactions, more or less individually, were studied earlier, Edward Haskell (1949) gave an integrative approach to the thematic, proposing a classification of "co-actions", later adopted by biologists as "interactions". Close and long-term interactions are described as

; symbioses that are mutually beneficial are called mutualistic.

Short-term interactions

Short-term interactions, including

and

, are extremely important in

and

. These are short-lived in terms of the duration of a single interaction: a predator kills and eats a prey; a pollinator transfers pollen from one flower to another; but they are extremely durable in terms of their influence on the evolution of both partners. As a result, the partners coevolve.

Predation

In predation, one organism, the predator, kills and eats another organism, its prey. Predators are adapted and often highly specialized for hunting, with acute senses such as vision,

hearing Hearing, or auditory perception, is the ability to perceive sounds through an organ, such as an ear, by detecting vibrations as periodic changes in the pressure of a surrounding medium. The academic field concerned with hearing is audit ...

, or smell. Many predatory animals, both

vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...

and

invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...

, have sharp

claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...

s or jaws to grip, kill, and cut up their prey. Other adaptations include stealth and

aggressive mimicry Aggressive mimicry is a form of mimicry in which predators, parasites, or parasitoids share similar signals, using a harmless model, allowing them to avoid being correctly identified by their prey or host. Zoologists have repeatedly compare ...

that improve hunting efficiency. Predation has a powerful selective effect on prey, causing them to develop

antipredator adaptation Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avo ...

s such as

warning coloration Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...

alarm call In animal communication, an alarm signal is an antipredator adaptation in the form of signals emitted by social animals in response to danger. Many primates and birds have elaborate alarm calls for warning conspecifics of approaching predato ...

s and other signals,

camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...

and defensive spines and chemicals. Predation has been a major driver of evolution since at least the

Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...

period. Over the last several decades, microbiologists have discovered a number of fascinating microbes that survive by their ability to prey upon others. Several of the best examples are members of the genera '' Daptobacter'' ( Campylobacterota), ''

Bdellovibrio ''Bdellovibrio'' is a genus of Gram-negative, obligate aerobic bacteria. One of the more notable characteristics of this genus is that members can prey upon other Gram-negative bacteria and feed on the biopolymers, e.g. proteins and nucleic ac ...

'', and ''

Vampirococcus ''Vampirococcus'' is an informally described genus of ovoid Gram-negative bacteria, but the exact phylogeny remains to be determined. This predatory prokaryote was first described in 1983 by Esteve et al. as small, anaerobic microbe about 0.6 � ...

''. Bdellovibrios are active hunters that are vigorously motile, swimming about looking for susceptible Gram-negative bacterial prey. Upon sensing such a cell, a bdellovibrio cell swims faster until it collides with the prey cell. It then bores a hole through the outer membrane of its prey and enters the periplasmic space. As it grows, it forms a long filament that eventually forms septae and produces progeny bacteria. Lysis of the prey cell releases new bdellovibrio cells. Bdellovibrios will not attack mammalian cells, and Gram-negative prey bacteria have never been observed to acquire resistance to bdellovibrios. This has raised interest in the use of these bacteria as a "probiotic" to treat infected wounds. Although this has not yet been tried, one can imagine that with the rise in antibiotic-resistant pathogens, such forms of treatments may be considered viable alternatives.

Pollination

In pollination, pollinators including

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

s (

entomophily Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, som ...

), some

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s ( ornithophily), and some bats, transfer

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...

from a male flower part to a female flower part, enabling

fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...

, in return for a reward of pollen or nectar. The partners have coevolved through geological time; in the case of insects and

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

s, the coevolution has continued for over 100 million years. Insect-pollinated flowers are adapted with shaped structures, bright colours, patterns, scent, nectar, and sticky pollen to attract insects, guide them to pick up and deposit pollen, and reward them for the service. Pollinator insects like

bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...

s are adapted to detect flowers by colour, pattern, and scent, to collect and transport pollen (such as with bristles shaped to form pollen baskets on their hind legs), and to collect and process nectar (in the case of

honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...

s, making and storing

honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...

). The adaptations on each side of the interaction match the adaptations on the other side, and have been shaped by

natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...

on their effectiveness of pollination.

Seed dispersal

Seed dispersal is the movement, spread or transport of

seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...

s away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their propagules, including both abiotic vectors such as the wind and living ( biotic) vectors like birds. Seeds can be dispersed away from the parent plant individually or collectively, as well as dispersed in both space and time. The patterns of seed dispersal are determined in large part by the dispersal mechanism and this has important implications for the demographic and genetic structure of plant populations, as well as migration patterns and species interactions. There are five main modes of seed dispersal:

gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the stro ...

, wind, ballistic, water, and by animals. Some plants are

serotinous Serotiny in botany simply means 'following' or 'later'. In the case of serotinous flowers, it means flowers which grow following the growth of leaves, or even more simply, flowering later in the season than is customary with allied species. Havi ...

and only disperse their seeds in response to an environmental stimulus. Dispersal involves the letting go or detachment of a diaspore from the main parent plant.

Symbiosis: long-term interactions

The six possible types of

are mutualism, commensalism, parasitism, neutralism, amensalism, and competition. These are distinguished by the degree of benefit or harm they cause to each partner.

Mutualism

Mutualism is an interaction between two or more species, where species derive a mutual benefit, for example an increased

carrying capacity The carrying capacity of an environment is the maximum population size of a biological species that can be sustained by that specific environment, given the food, habitat, water, and other resources available. The carrying capacity is defined as ...

. Similar interactions within a species are known as

co-operation Cooperation (written as co-operation in British English) is the process of groups of organisms working or acting together for common, mutual, or some underlying benefit, as opposed to working in competition for selfish benefit. Many animal ...

. Mutualism may be classified in terms of the closeness of association, the closest being symbiosis, which is often confused with mutualism. One or both species involved in the interaction may be

obligate {{wiktionary, obligate As an adjective, obligate means "by necessity" (antonym ''facultative'') and is used mainly in biology in phrases such as: * Obligate aerobe, an organism that cannot survive without oxygen * Obligate anaerobe, an organism tha ...

, meaning they cannot survive in the short or long term without the other species. Though mutualism has historically received less attention than other interactions such as predation,Begon, M., J.L. Harper and C.R. Townsend. 1996. ''Ecology: individuals, populations, and communities'', Third Edition. Blackwell Science Ltd., Cambridge, Massachusetts, USA. it is an important subject in ecology. Examples include

cleaning symbiosis Cleaning symbiosis is a mutually beneficial association between individuals of two species, where one (the cleaner) removes and eats parasites and other materials from the surface of the other (the client). Cleaning symbiosis is well-known amon ...

gut flora Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut m ...

, Müllerian mimicry, and

nitrogen fixation Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmo ...

by bacteria in the root nodules of

legumes A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock for ...

Commensalism

Commensalism benefits one organism and the other organism is neither benefited nor harmed. It occurs when one organism takes benefits by interacting with another organism by which the host organism is not affected. A good example is a

remora The remora (), sometimes called suckerfish, is any of a family (Echeneidae) of ray-finned fish in the order Carangiformes. Depending on species, they grow to long. Their distinctive first dorsal fins take the form of a modified oval, sucker-li ...

living with a

manatee Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living speci ...

. Remoras feed on the manatee's faeces. The manatee is not affected by this interaction, as the remora does not deplete the manatee's resources.

Parasitism

Parasitism is a relationship between species, where one organism, the

parasite 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 h ...

, lives on or in another organism, the host, causing it some harm, and is adapted structurally to this way of life. The parasite either feeds on the host, or, in the case of intestinal parasites, consumes some of its food.

Neutralism

Neutralism (a term introduced by

Eugene Odum Eugene Pleasants Odum (September 17, 1913 – August 10, 2002) was an American biologist at the University of Georgia known for his pioneering work on ecosystem ecology. He and his brother Howard T. Odum wrote the popular ecology textbook, ''Fun ...

) describes the relationship between two species that interact but do not affect each other. Examples of true neutralism are virtually impossible to prove; the term is in practice used to describe situations where interactions are negligible or insignificant.

Amensalism

Amensalism (a term introduced by Haskell) is an interaction where an organism inflicts harm to another organism without any costs or benefits received by itself. Amensalism describes the adverse effect that one organism has on another organism (figure 32.1). This is a unidirectional process based on the release of a specific compound by one organism that has a negative effect on another. A classic example of amensalism is the microbial production of antibiotics that can inhibit or kill other, susceptible microorganisms. A clear case of amensalism is where sheep or cattle trample grass. Whilst the presence of the grass causes negligible detrimental effects to the animal's hoof, the grass suffers from being crushed. Amensalism is often used to describe strongly asymmetrical competitive interactions, such as has been observed between the Spanish ibex and weevils of the genus ''Timarcha'' which feed upon the same type of shrub. Whilst the presence of the weevil has almost no influence on food availability, the presence of ibex has an enormous detrimental effect on weevil numbers, as they consume significant quantities of plant matter and incidentally ingest the weevils upon it. Amensalisms can be quite complex. Attine ants (ants belonging to a New World tribe) are able to take advantage of an interaction between an actinomycete and a parasitic fungus in the genus '' Escovopsis''. This amensalistic relationship enables the ant to maintain a mutualism with members of another fungal genus, ''Leucocoprinus''. Amazingly, these ants cultivate a garden of ''Leucocoprinus'' fungi for their own nourishment. To prevent the parasitic fungus ''Escovopsis'' from decimating their fungal garden, the ants also promote the growth of an actinomycete of the genus ''Pseudonocardia'', which produces an antimicrobial compound that inhibits the growth of the ''Escovopsis'' fungi.

Competition

Competition can be defined as an interaction between

s or species, in which the fitness of one is lowered by the presence of another. Competition is often for a resource such as

food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is in ...

water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as ...

, or

territory A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...

in limited supply, or for access to females for reproduction. Competition among members of the same species is known as intraspecific competition, while competition between individuals of different species is known as

interspecific competition Interspecific competition, in ecology, is a form of competition in which individuals of ''different'' species compete for the same resources in an ecosystem (e.g. food or living space). This can be contrasted with mutualism, a type of symbiosis. ...

. According to the

competitive exclusion principle In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the sligh ...

, species less suited to compete for resources should either adapt or die out. According to

evolutionary theory Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

, this competition within and between species for resources plays a critical role in

Non-trophic interactions

Foundation species enhance food web complexity

Some examples of non-trophic interactions are habitat modification, mutualism and competition for space. It has been suggested recently that non-trophic interactions can indirectly affect food web topology and

trophic dynamics A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...

by affecting the species in the network and the strength of trophic links. A number of recent theoretical studies have emphasized the need to integrate trophic and non-trophic interactions in ecological network analyses. The few empirical studies that address this suggest food web structures (network topologies) can be strongly influenced by species interactions outside the trophic network. However these studies include only a limited number of coastal systems, and it remains unclear to what extent these findings can be generalized. Whether non-trophic interactions typically affect specific species, trophic levels, or functional groups within the food web, or, alternatively, indiscriminately mediate species and their trophic interactions throughout the network has yet to be resolved. Some studies suggest sessile species with generally low trophic levels seem to benefit more than others from non-trophic facilitation, while other studies suggest facilitation benefits higher trophic and more mobile species as well. A 2018 study by Borst ''et al.''. tested the general hypothesis that foundation species — spatially dominant habitat-structuring organisms Angelini C, Altieri AH, Silliman BR, Bertness MD. Interactions among foundation species and their consequences for community organization, biodiversity, and conservation. Bioscience. 2011;61(10):782–9. – modify food webs by enhancing their size as indicated by species number, and their complexity as indicated by link density, via facilitation of species, regardless of ecosystem type (see diagram). Additionally, they tested that any change in food web properties caused by foundation species occurs via random facilitation of species throughout the entire food web or via targeted facilitation of specific species that belong to certain

trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it ...

s or functional groups. It was found that species at the base of the food web are less strongly, and carnivores are more strongly facilitated in foundation species' food webs than predicted based on random facilitation, resulting in a higher mean trophic level and a longer average chain length. This indicates foundation species strongly enhance food web complexity through non-trophic facilitation of species across the entire trophic network. Although foundation species are part of the food web like any other species (e.g. as prey or predator), numerous studies have shown that they strongly facilitate the associated community by creating new habitat and alleviating physical stress. This form of non-trophic facilitation by foundation species has been found to occur across a wide range of ecosystems and environmental conditions. In harsh coastal zones, corals, kelps, mussels, oysters, seagrasses, mangroves, and salt marsh plants facilitate organisms by attenuating currents and waves, providing aboveground structure for shelter and attachment, concentrating nutrients, and/or reducing desiccation stress during low tide exposure. In more benign systems, foundation species such as the trees in a forest, shrubs and grasses in savannahs, and macrophytes in freshwater systems, have also been found to play a major habitat-structuring role. Ultimately, all foundation species increase habitat complexity and availability, thereby partitioning and enhancing the niche space available to other species.