Pheromone

A pheromone (from Ancient Greek φέρω (phérō) 'to bear' and hormone) is a chemical that is secreted or excreted by an organism, which triggers a social response in members of the same species. There are alarm pheromones, food trail pheromones, sex pheromones, and many others that affect behavior or physiology. Pheromones are used by many organisms, from basic unicellular prokaryotes to complex multicellular eukaryotes. Their use among insects has been particularly well documented. In addition, some vertebrates, plants and ciliates communicate by using pheromones. The ecological functions and evolution of pheromones are a major topic of research in the field of chemical ecology.

Background The portmanteau word "pheromone" was coined by Peter Karlson and Martin Lüscher in 1959, based on the Greek φέρω phérō ('I carry') and ὁρμων hórmōn ('stimulating'). Pheromones are also sometimes classified as ecto-hormones ("ecto-" meaning "outside"). They were researched earlier by various scientists, including Jean-Henri Fabre, Joseph A. Lintner, Adolf Butenandt, and ethologist Karl von Frisch who called them various names, such as "alarm substances". These chemical messengers are transported outside of the body and affect neurocircuits, including the autonomous nervous system with hormone or cytokine mediated physiological changes, inflammatory signaling, immune system changes and/or behavioral change in the recipient. They proposed the term to describe chemical signals from conspecifics that elicit innate behaviors soon after the German biochemist Adolf Butenandt had characterized the first such chemical, bombykol, a chemically well-characterized pheromone released by the female silkworm to attract mates.

Categorization by function

Aggregation

Aggregation pheromones function in mate choice, overcoming host resistance by mass attack, and defense against predators. A group of individuals at one location is referred to as an aggregation, whether consisting of one sex or both sexes. Male-produced sex attractants have been called aggregation pheromones, because they usually result in the arrival of both sexes at a calling site and increase the density of conspecifics surrounding the pheromone source. Most sex pheromones are produced by the females; only a small percentage of sex attractants are produced by males. Aggregation pheromones have been found in members of the Coleoptera, Collembola, Diptera, Hemiptera, Dictyoptera, and Orthoptera. In recent decades, aggregation pheromones have proven useful in the management of many pests, such as the boll weevil (Anthonomus grandis), the pea and bean weevil (Sitona lineatus), and stored product weevils (e.g. Sitophilus zeamais, Sitophilus granarius, and Sitophilus oryzae). Aggregation pheromones are among the most ecologically selective pest suppression methods. They are non-toxic and effective at very low concentrations.

Alarm

Some species release a volatile substance when attacked by a predator that can trigger flight (in aphids) or aggression (in ants, bees, termites, and wasps) in members of the same species. For example, Vespula squamosa use alarm pheromones to alert others to a threat. In Polistes exclamans, alarm pheromones are also used as an alert to incoming predators. Pheromones also exist in plants: Certain plants emit alarm pheromones when grazed upon, resulting in tannin production in neighboring plants. These tannins make the plants less appetizing to herbivores. An alarm pheromone has been documented in a mammalian species. Alarmed pronghorn, Antilocapra americana flair their white rump hair and exposes two highly odoriferous glands that releases a compound described having the odor "reminiscent of buttered popcorn". This sends a message to other pronghorns by both sight and smell about a present danger. This scent has been observed by humans 20 to 30 meters downwind from alarmed animals. The major odour compound identified from this gland is 2-pyrrolidinone.

Epideictic Epideictic pheromones are different from territory pheromones, when it comes to insects. Fabre observed and noted how "females who lay their eggs in these fruits deposit these mysterious substances in the vicinity of their clutch to signal to other females of the same species they should clutch elsewhere." It may be helpful to note that the word epideictic, having to do with display or show (from the Greek 'deixis'), has a different but related meaning in rhetoric, the human art of persuasion by means of words.

Territorial

Laid down in the environment, territorial pheromones mark the boundaries and identity of an organism's territory. Cats and dogs deposit these pheromones by urinating on landmarks that mark the perimeter of the claimed territory. In social seabirds, the preen gland is used to mark nests, nuptial gifts, and territory boundaries with behavior formerly described as 'displacement activity'.

Trail

Social insects commonly use trail pheromones. For example, ants mark their paths with pheromones consisting of volatile hydrocarbons. Certain ants lay down an initial trail of pheromones as they return to the nest with food. This trail attracts other ants and serves as a guide. As long as the food source remains available, visiting ants will continuously renew the pheromone trail. The pheromone requires continuous renewal because it evaporates quickly. When the food supply begins to dwindle, the trail-making ceases. Pharaoh ants (Monomorium pharaonis) mark trails that no longer lead to food with a repellent pheromone, which causes avoidance behaviour in ants. Repellent trail markers may help ants to undertake more efficient collective exploration. The army ant Eciton burchellii provides an example of using pheromones to mark and maintain foraging paths. When species of wasps such as Polybia sericea found new nests, they use pheromones to lead the rest of the colony to the new nesting site. Gregarious caterpillars, such as the forest tent caterpillar, lay down pheromone trails that are used to achieve group movement.

Sex

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