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Northern gannet

The northern gannet (Morus bassanus) is a seabird, the largest species of the gannet family, Sulidae. It is native to the coasts of the Atlantic Ocean, breeding in Western Europe and Northeastern North America. It is the largest seabird in the northern Atlantic. The sexes are similar in appearance. The adult northern gannet has a mainly white streamlined body with a long neck, and long and slender wings. It is 87 – long with a 170 – wingspan. The head and nape have a buff tinge that is more prominent in breeding season, and the wings are edged with dark brown-black feathers. The long, pointed bill is blue-grey, contrasting with black, bare skin around the mouth and eyes. Juveniles are mostly grey-brown, becoming increasingly white in the five years it takes them to reach maturity.

Nesting takes place in colonies on both sides of the North Atlantic, the largest of which are at Bass Rock (75,000 pairs as of 2014), St. Kilda (60,000 pairs as of 2013) and Ailsa Craig (33,000 pairs as of 2014) in Scotland, Grassholm in Wales, and Bonaventure Island (60,000 pairs in 2009) off the coast of Quebec. Its breeding range has extended northward and eastward, with colonies being established on Russia's Kola Peninsula in 1995 and Bear Island (the southernmost island of Svalbard), in 2011. Colonies are mostly located on offshore islands with cliffs, from which the birds can more easily launch into the air. The northern gannet undertakes seasonal migrations and catches fish (which are the mainstay of its diet) by making high-speed dives into the sea.

The northern gannet was previously hunted for food in certain parts of its range, and although that practice still continues in the Outer Hebrides of Scotland and the Faroe Islands, the bird faces few other natural or man-made threats. Since its population is growing, the International Union for Conservation of Nature considers it a least-concern species. Because it is both a conspicuous and a common bird, it is referred to in several ancient myths and legends.

Taxonomy
The Swiss naturalist Conrad Gessner gave the northern gannet the name Anser bassanus or scoticus in the 16th century, and noted that the Scots called it a solendguse. The former name was also used by the English naturalist Francis Willughby in the 17th century; the species was known to him from a colony in the Firth of Forth and from a stray bird that was found near Coleshill, Warwickshire. It was one of the many species originally described by the Swedish zoologist Carl Linnaeus in the landmark 1758 10th edition of his Systema Naturae, where it was given the binomial name Pelecanus bassanus. The French biologist Brisson placed it in the genus Sula in 1760, and his compatriot Louis Pierre Vieillot moved the species to his new genus Morus in 1816. Morus is derived from Ancient Greek moros, meaning "foolish", and refers to the lack of fear shown by breeding gannets and boobies, which enables them to be easily killed. The specific name bassanus is from the Bass Rock in the Firth of Forth. The ornithologist Bryan Nelson in 1978 supported the species' inclusion in Sula as he felt the differences in anatomy, behaviour, ecology and morphology between gannets and boobies were not sufficient to warrant separate genera.

Charles Lucien Bonaparte described the American populations as Sula americana in 1838, though the basis for distinguishing them from the European species was unclear and the name is now considered to be a synonym.

"Northern gannet" has been designated as the official common name for the species by the International Ornithologists' Union (IOC). It is also known as the North Atlantic gannet. Gannet is derived from Old English ganot, meaning "strong or masculine", which is ultimately from the same Old Germanic root as gander. Soland goose and similar old names for the northern gannet such as solan or solan goose derive from a hypothetical Scottish Gaelic sulan, itself borrowed from the Old Norse sula. The literal meaning is "cleft stick", referring to the appearance of the conspicuous crossed black wing tips on a perched northern gannet. Old regional names such as Norfolk's "herring gant" or Yorkshire's "mackerel gant" refer to typical fish prey. Lincolnshire's gaunt, although derived from the same Germanic root, usually applies to the great crested grebe, but the English writer Richard Hakluyt used the term in 1600 to refer to the gannet, "a great White foule". Young birds have been called "spotted booby" or "parliament goose", the former term referring to their plumage. The feeding habits of the gannet have led to its name being used as slang for a gluttonous person, a usage first recorded in 1929.

The Sulidae, the gannets and boobies, appeared about 30 million years ago. Early Sulidae fossils resembled the boobies, although they were more aquatic, the gannets splitting off later, about 16 million years ago. The gannets evolved in the northern hemisphere, later colonising the southern oceans. The most ancient extant species may be the Abbott's booby, possibly the sole survivor of an otherwise extinct separate lineage. A 2011 genetic study of nuclear and mitochondrial DNA suggests that the ancestor of the gannets arose around 2.5 million years ago before splitting into northern and southern lineages. The latter then split into the Cape and Australasian gannets around 0.5 million years ago. The three gannets are generally considered to be separate species forming a superspecies, though they have also formerly been classified as subspecies of Sula bassanus.

Description
An adult northern gannet has a 170 – wingspan, and is 87 – long and weighs 2.3 –, making it the largest gannet and the largest seabird native to the western Palearctic. The two sexes are generally of a similar size and appearance. The plumage is white with dark brown to black wing tips; the primary flight feathers, primary coverts and alulae are dark. The head and neck are tinged buff-yellow, becoming much more prominent in the breeding season. Males are more deeply coloured than females. The eyes are surrounded by black bare skin and have a cobalt blue orbital ring, the iris is light blue-grey with a fine dark outer ring. In birds that survive avian influenza, the irises become black.

The beak is long, strong and conical with a slight downcurve at the end and a sharp cutting edge. In adults, the beak is blue-grey with dark grey or black edges. There is a black groove running the length of the mandible that merges into the skin around the eyes. A black band of bare skin also separates the pale feathers of the forehead and throat from the bill, which gives the gannet its distinctive face markings. The four-toed feet are joined by a membrane that can vary in colour from dark grey to dark brown. There are coloured lines running along the toes that continue along up the legs. These are typically greenish-yellow in males and bluish in females and probably have a role in mating. Fledglings are dark grey to slate-grey with upperparts and wings finely speckled with white. There is a prominent V-shaped white area under the rump. The wing tips and tail are dark brown-black, partly tipped with white. The bill and iris are dark brown. They can weigh more than 4 kg by the time they leave the nest at about 10 weeks of age. In the second year, the bird's appearance changes depending on the different phases of moulting: they can have adult plumage at the front and continue to be brown at the rear. Gannets gradually acquire more white in subsequent seasons until they reach maturity after five years.

Northern gannets are slightly larger and thicker-billed than Cape or Australian gannets. The northern gannet has more white in the wings and an all-white tail, the other species having black tips to their tail feathers. Individuals on the west coast of Africa could be confused with vagrant masked boobies, though the latter is smaller overall, lacks the buff tinge to the head, and has a black tail. From a distance, or in poor visibility, albatrosses can be confused with northern gannets, particularly those with immature plumage that have more black on the wings.

Adaptations for diving
Northern gannets have streamlined bodies adapted for plunge-diving at high speed, including powerful neck muscles, and a spongy bone plate at the base of the bill. The nostrils are inside the bill and can be closed to prevent water entry; the eyes are protected by strong nictitating membranes. There are subcutaneous air sacs in the lower body and along the sides. Other air sacs are located between the sternum and the pectoral muscles and between the ribs and the intercostal muscles. These sacs are connected to the lungs and are filled with air when the bird breathes in. The air can be returned to the lungs by muscle contractions.

The feathers are waterproof, which allows the birds to spend long periods in water. A water-impermeable secretion produced by a sebaceous gland covers the feathers and the birds spread it across their body using their beak or head. Individuals have a subcutaneous fat layer, dense down feathers and tightly overlapping feathers that help them withstand low temperatures. A reduced blood flow in the webbing on their feet outside of the breeding season also helps to maintain body temperature when the birds swim.

Call

The northern gannet is a loud and vocal bird, particularly in the colony. Its typical call is a harsh arrah-arrah or urrah-urrah, which is emitted upon arriving or when challenging other gannets at the colony. The call is shortened to a rah rah when fishing or collecting nesting material, and lengthened to a ooo-ah when taking off. The calls of the sexes are similar. According to Nelson northern gannets can recognize the call of their breeding partner, their chicks and birds in neighbouring nests. Individuals from outside this sphere are treated with more aggression.

Distribution and habitat
The northern gannet's breeding range is on both sides of the North Atlantic on coasts influenced by the Gulf Stream, There are colonies in the Gulf of Saint Lawrence and on the islands off the east coast of Canada. They normally nest in large colonies, on cliffs overlooking the ocean or on small rocky islands. The water needs to be cool enough for Atlantic mackerel and herring, which are the main food source for the northern gannet. These areas also overlie the continental shelf. Northern gannet colonies can be found in the far north in regions that are very cold and stormy, and Nelson has suggested that they can survive in these regions for several reasons, including the combination of body weight and a powerful beak that allows them to capture strong muscular fish, and the ability to dive to great depths and capture prey far from the cliffs. Their fat reserves act as weight when diving and as reserves during extended periods without food.

The northern limit of their breeding area depends on the presence of waters that are free of sea ice during the breeding season. Therefore, while Greenland and Svalbard offer suitable breeding sites, the Arctic regions have summers that are too short to allow the northern gannets to lay their eggs and raise a brood, which requires between 26 and 30 weeks. The southern limit of their distribution mainly depends on the presence of sufficient prey. There is fossil evidence of northern gannets breeding on Crete in the Pleistocene.

Breeding colonies
Some northern gannet breeding colonies have been recorded as being located in the same place for hundreds of years. The cliffs containing the colonies appear white when seen from a distance, due to the number of nesting birds present on them. There is a written record of a colony on the island of Lundy from 1274. There were only 70 nests by 1871, and the colony finally disappeared by 1909 at the latest. More than two-thirds of the world population breeds around the coasts of the British Isles. Colonies include:

 * Scotland
 * Bass Rock, off the Firth of Forth, first recorded in 1493. In 2004, it contained more than 48,000 nests. By 2014 this had increased to over 75,000, making it the largest colony in the world.
 * Northern gannets began a colony at Troup Head in Aberdeenshire in 1988, and by 2014 it held an estimated 6,456 pairs.
 * Saint Kilda and Sula Sgeir, in the Hebrides, Scotland. The former colony was estimated at 60,290 pairs and the latter 11,230 pairs in 2013. A colony on the Flannan Isles almost doubled in size to 5,280 pairs in 2013. Further south, the island of Ailsa Craig in the Firth of Clyde hosted an estimated 33,226 pairs in 2014. Ailsa Craig has been known as a colony since 1583.
 * In Shetland, there were an estimated 25,580 breeding pairs at Hermaness, 11,786 on Noss, and 3,591 on Fair Isle in 2013, while Orkney had an estimated 4,550 pairs at Sule Stack. Sule Skerry's breeding population rose from 57 to 1,870 pairs between 2003 and 2013.

 * Wales
 * Grassholm off the Pembrokeshire coast in Wales had an estimated 36,011 pairs in 2015. Gannets began nesting here between 1820 and 1860, though never in great numbers, only exceeding 300 pairs by 1913.

 * England
 * Bempton Cliffs hosted an estimated 12,494 pairs in 2015, more than tripling in size in 12 years.

 * Ireland
 * Little Skellig, a small island located about 12 km off the Iveragh Peninsula of County Kerry is the largest colony in Ireland, hosting around 30,000 breeding pairs. Known as a gannetry before 1700, human impact had reduced the population to 30 pairs by 1880 before rapidly increasing to around 10,000 pairs by 1906, the fastest recovery ever recorded at a gannet colony. There are small colonies on Ireland's southern coast, on Bull Rock, County Cork, Clare Island, County Mayo and Great Saltee Island, County Wexford.

 * Alderney. Around 7,500 pairs nest on two islets—Ortac and Les Etacs—off the coast of Alderney in the English Channel, the first birds only nesting there in the 1940s. These birds migrate further south to the Mediterranean and west coast of Africa than more northerly breeding colonies.
 * France. The French island of Rouzic in the Jentilez archipelago off the coast of Brittany hosts the southernmost breeding colony of northern gannets. Established in the late 1930s, it had grown to over 11,500 breeding pairs by 1995. Pairs have nested sporadically with varying success along the Mediterranean coast.
 * Germany. Northern gannets have reappeared on Heligoland in 1991 and around 680 breeding pairs have been counted in 2016.
 * Italy. A pair raised a chick successfully for three successive seasons from 2013 to 2015 on a moored boat at Porto Venere in northern Italy, after birds had been seen there since 1993.
 * The Faroe Islands. Mykines in the Faroe Islands held about 2,000 breeding pairs in 2000, although that had risen to 2,500 pairs by 2014.
 * Iceland. Eldey, a small island located about 16 km off the coast of the Reykjanesskagi Peninsula, Iceland, hosting around 16,300 breeding pairs in 1962, and a similar number in 2008. Iceland has several small colonies along its coast, and on Grimsey, around 40 km north.
 * Norway. As of 2016, there are an estimated 6,900 breeding pairs in Norway. Northern gannets first bred in the south of the country in the mid 1940s at Runde in Møre og Romsdal. In 1967 a colony was established at Syltefjord, within the Arctic Circle at 70°N. Colonies were also established on low islets in Lofoten and Vesterålen, but declined due to increasing numbers of white-tailed eagles. Once rare, these were protected in 1968 and their numbers have risen. It is unclear why the eagles have not impacted on gannet numbers further north. In 2011, gannets began breeding on Bear Island, southernmost island of Svalbard, likely due to the warming of the Barents Sea and influx of fish. The colony numbered 52 pairs in 2016.
 * Russia. There are around 250 pairs in a colony at Kharlov on the Kola Peninsula in Russia. Established in 1995, this is also thought to be due to the warming of the Barents Sea.
 * North America. There are six breeding colonies along the coast of Canada. The Gulf of Saint Lawrence hosts three colonies—Bonaventure Island, Bird Rocks and Anticosti Island—and there are three off the eastern and southeastern coast of Newfoundland (Cape St. Mary's Ecological Reserve, Funk Island, Baccalieu Island Ecological Reserve. Bonaventure Island off the south coast of the Gaspé Peninsula of Quebec is the largest colony with 60,000 nests as of 2009.

Migration
After the breeding season, adult northern gannets disperse over a wide area although they travel no more than 800 to 1600 km from the breeding colony. It is not known if all birds from one colony migrate to the same over-wintering area. Many adults migrate to the west of the Mediterranean, passing over the Strait of Gibraltar and flying over land as little as possible. Other birds follow Africa's Atlantic coastline to arrive in the Gulf of Guinea. Immature northern gannets from colonies in Canada fly to the Gulf of Mexico, much further south than the adults. The immature gannets migrate southwards for great distances and have been recorded as far south as Ecuador. In their second year some birds return to the colony they were born in, where they arrive later than the mature birds. They then migrate south again at the end of the breeding season, but travel shorter distances in this second migration. Gannets from Alderney have been tracked since 2015 to gain better knowledge of their movements. One individual was found to have travelled from its colony in Alderney to Scandinavian waters, a round trip of around 2,700 km.

The species has been recorded as a vagrant in many central and eastern European countries as far south and west as the Black Sea, and also in Bermuda, Cuba, Cyprus, Egypt, Kazakhstan, Jan Mayen and Syria. In February 2016, one was recorded from Ceará in northeastern Brazil—the first sighting in the Southern Hemisphere.

Behaviour
The wings of the northern gannet are long and narrow and are positioned towards the front of the body, allowing efficient use of air currents when flying. Even in calm weather they can attain velocities of between 55 and 65 km/h although their flying muscles are relatively small: in other birds flying muscles make up around 20% of total weight, while in northern gannets the flying muscles are less than 13%. Despite their speed, they cannot manoeuvre in flight as well as other seabirds. Northern gannets need to warm up before flying. They also walk with difficulty and this means that they have problems getting airborne from a flat area. They take off from water by facing into the wind and strongly beating their wings. In light winds and high waves they are sometimes unable to take off and they can become beached.

Northern gannets alight on land using angled wings, fanned tail and raised feet to control their speed, not always successfully, since damaged or broken wings were recorded as a frequent cause of death in adults at one colony.

Feeding
Northern gannets forage for food during the day, generally by diving at high speed into the sea. They search for food both near to their nesting sites but also further out to sea. Birds that are feeding young have been recorded searching for food up to 320 km from their nest. It has been found that 2% of birds nesting in the colony on Bass Rock search for fish at Dogger Bank, between 280 and 320 km away. It is likely that they fly further than this while foraging, possibly up to double the distance; normally they fly less than 150 km. Some studies have found that the duration and direction of flights made while foraging for food are similar for both sexes, although there are significant differences in the search behaviour of males and females. Female northern gannets are not only more selective than males in choosing a search area: they also make longer and deeper dives and spend more time floating on the surface than males. Gannets will follow fishing boats or cetaceans to find discarded or injured fish. They forage from heights of up to 70 m with no clear preference, and typically dive between 11 and 60 m. They dive with their bodies straight and rigid, wings tucked close to the body but angled back, extending beyond the tail, before piercing the water like an arrow. They control the direction of the dive using their wings and tail, and fold their wings against the body just before impact. Birds can hit the water at speeds of up to 100 km/h. This allows them to penetrate up to 11 m below the surface, and they will swim down to an average 19.7 m, sometimes deeper than 25 m. The bird's subcutaneous air sacs may have a role in controlling their buoyancy.

Gannets usually push their prey deeper into the water and capture it as they return to the surface. When a dive is successful, they swallow the fish underwater before surfacing, and never fly with the fish in their bill. Larger fish are swallowed headfirst, smaller fish are swallowed sideways or tail-first. The fish is stored in a branched bag in the throat and does not cause drag when in flight. Their white colour helps other gannets to identify one of their kind and they can deduce the presence of a shoal of fish by this diving behaviour; this in turn facilitates group foraging, which makes capturing their prey easier. The colour also makes the gannet less visible to the fish underneath. Northern gannets also forage for fish while swimming with their head under water.

They eat mainly fish 2.5 – in length that shoal near the surface. Virtually any small fish (roughly 80–90% of their diet) or other small pelagic species (largely squid) will be taken opportunistically. Sardines, anchovies, haddock, smelt, Atlantic cod and other shoal-forming species are also eaten. M. bassanus takes huge quantities from the waters around Newfoundland for the energy requirements of their growing chicks. Given the economic importance of the prey taken, whether they are in competition with human fishing industry in the area is an important question which remains unassessed. (But given the small amount taken, NL's cod take is probably unaffected.) On the other hand, they profit off of discarded bycatch and processed parts   and the reduction in competition if humans are taking predatory fish.

M. bassanus inflicts significant wastage on the northwest Atlantic Salmo salar fishery, but does not actually eat much of them. This makes it difficult to resolve this conflict with the fishing industry in the area, as the only option would be to exterminate the birds, which is ecologically unacceptable.

Breeding
The oldest birds are the first to return to the northern gannet's breeding colonies. Birds not of breeding age arrive a few weeks later. In general, birds first return to a colony (not uncommonly the one in which they were hatched) when they are two or three years old. Once an individual has successfully bred in a colony it will not normally change to another. Nesting starts in March or April.

Immature birds stay on the edges of the colony. They may even make a nest but they do not breed until they are 4 or 5 years old. Some birds of this age occupy empty nests that they will aggressively defend if they have sat on them for two or three days. If an apparently empty nest has an owner, the immature bird will leave without a struggle when the owner arrives to take possession.

The preferred nesting sites are on coastal hillsides or cliffs. If these are not available northern gannets will nest in groups on islands or flat surfaces. As they find it more difficult to take off from such locations they will often cross the area occupied by an adjacent nest causing an aggressive reaction from the sitting pair; this means that the stress levels are higher in this type of colony than in those on steeper surfaces. Notwithstanding this, nests are always built close together and otherwise ideal nesting sites will not be used if they are some distance from a colony. On average there are 2.3 nests per square metre (1.9 per square yard). Both sexes fiercely defend the area around their nest. Where space allows, the distance between nests is double the reach of an individual. Nests are made from seaweed, plants, earth and debris from the sea. The males usually collect the materials. Nests are compact cups typically 30 – in height. The area which a nest occupies grows throughout the breeding season as the breeding pairs throw their excrement outside the nest. Over years, nests can reach 2 m in height.

Northern gannets lay one egg that on average weighs 104.5 g, which is light for such a large seabird. The egg is around 79 mm long by 50 mm wide and the shell is pale blue and translucent initially before fading to a chalky white surface that is easily stained. M. bassanus was among those birds most affected by the shell-thinning effect of DDT, which contributed in large degree to its ban. Where two eggs are found in a nest this is the result of two females laying an egg in the same nest or one egg being stolen from another nest. Northern gannets will lay a replacement egg if the first is lost. Incubation takes 42 to 46 days, during which time the egg is surrounded by the brooding bird's warm, webbed feet. Just before hatching begins, the brooding bird releases the egg from its feet to prevent the egg from breaking under the adult's weight as the chick breaks it open. This is a frequent cause of death for chicks of birds that are breeding for the first time. The process of breaking the eggshell can take up to 36 hours. The webbed feet are also used to cover the chicks, which are only rarely left alone by their parents. Chicks that are left unattended are often attacked and killed by other northern gannets. Newly hatched chicks are featherless and are dark blue or black in colour. In the second week of life they are covered in white down, replaced over the next five weeks by dark brown feathers flecked with white. Young chicks are fed regurgitated semi-digested fish by their parents, who open their mouths wide for their young to fetch the food from the back of their throats. Older chicks receive whole fish. Unlike the chicks of other species, northern gannet chicks do not move about the nest or flap their wings to ask for food: this reduces the likelihood that they will fall from the nest.

The adults feed their offspring for around 13 weeks, right up until the time they leave. The young birds fledge between 84 and 97 days old, departing by launching themselves off a cliff and flying—a procedure for which it is impossible to practice beforehand. If they leave the nest in bad weather they can be mortally wounded as they can be blown against the rocks. The young birds are attacked by adults if unattended. Once they leave the nest they stay at sea learning to fish and fly, their flight skills being too poor for them to return to the breeding ledges.

Northern gannets have only one brood a year. The survival rate for young birds for their first four years is 30% and the annual survival rate for adults is 91.9%. The typical lifespan after becoming adult is 17 years, and the maximum known age is 37 years 4 months 16 days. Gannet pairs are monogamous and may remain together over several seasons, if not for all of their lives. The pairs separate when their chicks leave the nest but they bond again the following year. Should one of the pair die, the other bird will find another mate.

Displays
Northern gannets exhibit many types of aggressive behaviour while they are nesting. Confrontations normally only take place between birds of the same sex. Females will lower their heads before an aggressive male that is defending its nest: this will expose the back of the female's neck and the male will take it in its beak and expel the female from the nest. A female will not react if a male approaches a nest but it will react fiercely if another female approaches. The fights between males occupying nests for the first time are particularly intense. Such fights can last for up to two hours and lead to serious injuries. Birds lunge at each other and lock bills, wrestling for extended periods while neighbours peck at them. The fights are preceded by threatening gestures, which are also seen outside the breeding season. Males demonstrate ownership of a nest by gesturing towards their neighbours in a bowing display; their heads and beaks point down, and their wings are held up and away from the body, yet folded backwards. The male moves his head from side to side before bowing forwards.

Males try to attract an available female after establishing a territory. The females will fly over the colony several times before landing. Their posture, with the neck stretched out, tells the male that they are available for courtship. The male will then shake their heads in a similar way to when they are guarding their nest, but with their wings closed. Mated pairs engage in a fencing display when the male arrives back at the nest. The two birds stand breast to breast with wings spread and bills extended vertically. They fence and scissor with their bills rapidly, calling loudly at the same time. Fencing is interspersed with bill bowing.

Predators and parasites
The northern gannet is not heavily predated. The only known habitual natural predators of adults are bald eagles, white-tailed eagles and golden eagles. Predators of eggs and nestlings include the great black-backed gull and American herring gull, common ravens, ermine, and red fox. Attacks at sea are insignificant, though large sharks and seals may rarely snatch a gannet out at sea.

Kleptoparasitism by skuas, particularly the great skua, occurs at breeding sites. The skua chases its victim until it disgorges its stomach contents, providing a meal for the attacker. Skuas may catch the tip of the gannet's wing, causing it to fall into the sea, or seize the tail to tip its victim into the water. The gannet is only released when it has regurgitated its catch.

External parasites include feather lice, although there are relatively few species and none are found on the head. As with grebes and divers, it may be that the short head feathers provide insufficient cover for the parasite. In one species, Michaelichus bassani, immature lice are found in the membranes lining the subcutaneous air-cells. Ixodes mites include the widespread I. uriae.

The spiny-headed worm Corynosoma tunitae appears to occur only in gannets and closely related seabird families such as the cormorants. The tapeworm Tetrabothrius bassani absorbs toxic heavy metals at a higher concentration than the gannet's own tissues, with an average 12 times as much cadmium as the gannet's pectoral muscles and 7–10 times the lead level of the bird's kidney and liver. Since levels of these toxic metals are detectable in the parasite earlier than in the host, the tapeworm might be used as an early indicator of marine pollution.

Conservation status
A 2004 survey counted 45 gannet breeding colonies and some 361,000 nests. The population is apparently growing between 3% and 5% a year, although this growth is concentrated in just a few colonies. Although northern gannet populations are now stable, their numbers were once greatly reduced due to loss of habitat, removal of eggs and killing of adults for their meat and feathers. In 1939, there were 22 colonies and some 83,000 nests, which means that the populations have increased fourfold since that time.

In 1992, the International Union for Conservation of Nature (IUCN) estimated the bird's population to be some 526,000. After taking into account an estimate produced for BirdLife International in 2004 of the European population, the IUCN revised its global population to between 950,000 and 1,200,000 individuals.

The IUCN lists northern gannets as a species of least concern, as they are widely distributed and as there is a large population that appears to be growing due to high breeding success, with 75% of eggs producing fledged young.

In culture
In Homer's Odyssey, the sea goddess Leucothea (Λευκοθέα "white goddess"), appears "in the likeness of a Gannet" and tells the shipwrecked Odysseus to discard his cloak and raft, instead offering him her veil to wind round himself which will save his life and enable him to reach land.

Another early reference to the gannet is in the 7th-century Old English epic poem The Seafarer.

An old myth from Mykines in the Faroe Islands tells of the giant Tórur seeking mercy following defeat at the hands of Óli, the islanders' head man and magician. In return, he gave them whales, driftwood logs and a bird unique to the archipelago, on condition that the inhabitants did not laugh at his gifts. Over time, the islanders forgot their promise, and lost the whales and logs, but fearful of losing a valuable food source, they never mocked the gannets that Tórur had given them.

Northern gannets have long been eaten for food. Birds, mainly the young, were taken from Bass Rock for at least 350 years until 1885, when the annual cull of about 1,500 individuals finally ceased, and Shetland gannets were sold as "Highland goose" in London restaurants during World War II. Views of the palatability of this bird are mixed, but as well as being a food for the poor it also regularly featured in Scottish royal banquets. In Scotland gannets were traditionally salted to preserve them until they got to market, this technique being replaced by partially cooking or smoking in the era of modern transport. They are normally served roasted, although sometimes raw when pickled or dried.

The best-known site was the remote island of St. Kilda, where adults and eggs were taken in the spring. The fat chicks, known locally as "gugas", were harvested from the precipitous cliffs in August, just before they could fly, and thrown to waiting boats far below. Much of the meat was salted in barrels for storage, but the rest of the bird was also used. Islanders paid their rent in feathers for stuffing pillows and furniture, the gannet stomachs were used to hold oil derived from the carcasses, and the breastbones served as lamp wicks.

Hunting on St. Kilda ceased in 1910, but the gannetry on Sula Sgeir is still exploited under a licence that permits 2,200 chicks to be taken each year. During the hunt, 10 men live on the island, and the cleaned birds are singed on a fire fuelled by their own oil-rich offal. The filleted birds are then taken to Stornoway, where each hunter receives 200 skins to give away or sell. The continuing existence of the practice of hunting and eating gannets attracts criticism in some quarters. The island's name "Sula Sgeir" itself derives from sula, meaning "gannet", and the Old Norse skerr, a skerry. Other sites that continued hunting into the 20th century were Eldey in Iceland, where the activity ceased in 1939, and Mykines, where small-scale culling still persists. About 500 young are culled for consumption each year in Mykines, using techniques similar to those of the Sula Sgeir hunts.

Although the Bass Rock population fell to fewer than 4,000 pairs in the early 19th century, the population soon recovered once hunting ceased, and St. Kilda was harvested sustainably for hundreds of years. Elsewhere, the recovery was less complete. The Bird Rocks colony in the Gulf of St Lawrence may once have held 250,000 birds, but unchecked hunting, including for fish bait, meant that the population was only 1,000 birds by 1932, despite government protection since 1904.

title: Northern gannet


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The term is relative and partially subjective, reflecting both objective qualities of a car, as well as the projected and perceived image of the vehicle marque. Luxury brands rank above premium brands, though there is no clear distinction between the two.

Traditionally, most luxury cars were large vehicles, though smaller sports-oriented models were produced. "Compact" luxury vehicles such as hatchbacks and off-road capable sport utility vehicles are more recent expansions of luxury qualities in various cars.

Increasing comfort features, materials quality, interior space, innovations, and or performance have consistently been competitive strategies between car makers practically throughout the history of the automobile.

Classification standards
Several car classification schemes include a luxury category, such as:
 * Australia: Since the year 2000, the Federal Government's luxury car tax applies to new vehicles over a certain purchase price, with higher thresholds applying for cars considered as fuel efficient. As of 2019, the thresholds were approximately AU$66,000 (US$,000) for normal cars and AU$76,000 (US$,000) for fuel-efficient cars.
 * Europe: Luxury cars are classified as F-segment vehicles in the European Commission classification scheme.
 * Italy: The term "auto di lusso" is used for luxury cars.
 * France: The term "voiture de luxe" is used for luxury cars.
 * Germany: The term Oberklasse (upper class) is used for luxury cars.
 * Russia: The term (автомобиль представительского класса ("representative class vehicle, also translated as luxury vehicle) is used for luxury cars.
 * Rental cars: The ACRISS Car Classification Code is a system used by many car rental companies to define equivalent vehicles across brands. This system includes "Luxury" and "Luxury Elite" categories (along with "Premium" and "Premium Elite" categories). The criteria for a vehicle to be considered "luxury" is not published.

Features
Luxury cars have traditionally emphasized higher levels of comfort and safety. Manufacturers often introduce new safety technologies and comfort amenities on luxury models before they are available on more affordable models. Some brands, like Audi and BMW have expanded their marketing by "introducing lesser priced and strip-down economy versions of their products."

Luxury vehicles can be a status symbol for conspicuous consumption. However, since many European luxury car buyers shy away from conspicuous consumption, brands offer buyers the option of removing exterior badges that identify the model name or engine size.

The suspension system of most luxury cars is tuned to prioritize ride quality over handling, however, some are marketed as "sports luxury" and have a greater emphasis on handling characteristics.

Layout and powertrain

Traditionally, luxury cars have used a front-engine, rear-wheel drive (FR) layout. The FR layout is more expensive to produce and produces lower fuel economy than a front-wheel drive layout; however, it allows for larger engines (particularly straight-six, V8, and V12) to be used.

Some American luxury cars during the 1970s through the 1990s switched to a front-wheel drive layout with transverse engine, due to the Arab Oil Embargo of 1973 and the 1979 fuel crises which eliminated many FR platforms in favor of the more economical front-wheel drive (FF) layout. From the early 2000s, several of these American luxury cars reverted to FR layouts.

Since the introduction of the Bentley Continental GT in 2003, additional luxury grand tourers feature all-wheel drive.

European manufacturers

Prior to World War II, a wide array of European producers made luxury cars, including Rolls-Royce, Bugatti, Delage, Delahaye, Talbot-Lago, Bentley, Alvis, Avions Voisin, Isotta Fraschini, Horch, Simson, Stoewer, Maybach, Mercedes-Benz, Hispano Suiza, Daimler Company, and Spyker.

France was a leading producer of powerful luxury automobiles before World War II. After World War II, the French government used puissance fiscale tax regulations to encourage manufacturers to build cars with small engines, and French motorists to buy them. The Maserati-powered Citroën SM and the Citroën C6 were arguably the last domestic French luxury cars. In the 2010s, some French manufacturers have attempted to develop luxury cars, however the lack of a historical legacy has hindered these efforts. In 2014, Citroën introduced DS Automobiles sub-brand to market luxury cars.

Pre World War II intermediate car manufacturers like Renault, Fiat, Opel, Lancia, Škoda, Riley, Praga, Peugeot, Hillman and Tatra made luxury cars but were forced to make economy cars and superminis post World War II. Following World War II, Germany rose to become an export powerhouse, building on success with the Mercedes-Benz brand, later joined by BMW, which acquired Rolls-Royce Motor Cars in 1998, as well as Volkswagen which much later controlled or acquired additional brands such as Audi, Porsche, Bentley, Lamborghini, and Bugatti brands.

In the Soviet Union, the manufacturer ZiL (then called Zis) began producing representational limousines in the mid-1930s. In the early 1950s, GAZ joined with the somewhat smaller "Chaika" model range. In 2018, Aurus Motors was established to produce luxury vehicles for the Russian market.

North American manufacturers

The luxury car phenomenon began at the start of the automobile industry when the wealthy frequently invested in manufacturing such models to gain social prestige. Emphasis was also placed on custom-built coachwork. The 1920s and 1930s were the apogee of production of these very large luxury automobiles from many manufacturers. The significant North American manufacturers from 1910 until 1940 included Auburn, Buick, Cadillac, Chrysler, Continental, Cord, Daniels, DeSoto, Duesenberg, Franklin, Imperial, LaFayette, LaSalle, Lincoln, Marmon, Packard, Peerless, Pierce Arrow, Ruxton, Stearns-Knight, and Stutz. The Great Depression put many luxury car manufacturers out of business; others would hold on before going defunct during the postwar era.

From 1946 until the late 1990s, Cadillac was the top-selling brand of luxury cars in the U.S., while Lincoln was second. The most successful and long-running model names during this era were the Cadillac DeVille, Lincoln Continental, and the Chrysler Imperial. The Lincoln Mark Series and Cadillac Eldorado were positioned in the personal luxury category, and competition between them continued into the 1990s.

The personal luxury car emerged into mass popularity and affordability as an America-specific category of popularly-priced cars made from the 1950s by the four domestic manufacturers (GM, Ford, Chrysler, and AMC) that reached peak popularity in the 1970s. The cars were stylized, mass-produced two-door coupés or convertibles, relying on standard components. These distinctively styled cars were targeting the needs of individual customers, not an entire family. The longest running model lines were the 1958-1997 Ford Thunderbird, 1956-1998 Lincoln Mark Series, and the 1967-2002 Cadillac Eldorado.

In 1990, American luxury brands dominated, with Cadillac selling over a quarter-million cars, and Lincoln had its best year ever at 231,660 units. However, the market was changing with an ever greater acceptance of smaller, more efficient imported luxury brands while at the same time, the domestic manufacturers were downsizing their models with product decisions that backfired on quality and brand respect.

Since the late 1990s, Japanese and German brands have sold the most luxury-type cars in the United States. However, the Cadillac Escalade has led the luxury SUV segment sales in the United States since its introduction in 1998, with the highest sales for 15 out of its first 20 years on the market.

In the 2000s, both Ford and General Motors produced luxury pickups: 2002-2013 Cadillac Escalade EXT, 2002-2003 Lincoln Blackwood, and 2006-2014 Lincoln Mark LT. In the late 2000s, the Cadillac CTS and Cadillac DTS led to a resurgence in the brand's luxury sedans. The equivalent sedan from the Ford group, the 2008 Lincoln MKS, was also regarded as a significant improvement over previous models. In 2010, BMW was the best-selling luxury vehicle manufacturer by sales, with Audi and Mercedes-Benz the second and third highest selling luxury brands.

East Asian manufacturers
Chinese manufacturer Hongqi was launched in 1958, making it the oldest Chinese luxury car marque. Later newcomers joined taking advantage of the rise of electric powertrains, with NEV brands such as Nio in 2014, Li Auto in 2015, HiPhi in 2019, and BYD in 2020 producing luxury electric and hybrid vehicles.

Japanese manufacturers have been producing luxury cars since the 1950s, including the Toyota Crown (1955–present), Prince/Nissan Gloria (1959–2004), Nissan Cedric (1960–2015), Mitsubishi Debonair (1964–1998), Nissan President (1965–2010), Toyota Century (1967–present), Mazda Luce/929 (1969–1991), and Honda Legend (1985–present).

Since the 1980s, overseas sales of Japanese luxury cars have increased, challenging traditional European luxury brands.

Several East Asian manufacturers have created sub-brands for the marketing of luxury cars. The first of these was the 1986 launch of Acura (a Honda sub-brand), followed by Lexus (Toyota) in 1989, Infiniti (Nissan) in 1989, and Genesis (Hyundai) in 2015.

Global financial crisis
The late-2000s global financial crisis was the first time since the Great Depression of the 1930s that the luxury car market suffered considerably, something not seen in previous economic downturns. Many such customers saw their net worth decline following the collapse in financial markets and real-estate values. For example, some of the steepest drop-offs came at the high end, including the BMW 7 Series and Rolls-Royce Phantom, and in 2010 Mercedes-Benz dropped the price of the W212 E-Class. The unusually sharp decline in luxury car sales has led observers to believe that there is a fundamental shift and reshaping of the luxury automotive market, with one industry official suggesting that the marques no longer command the premiums that they used to and another saying that conspicuous consumption was no longer attractive in poor economic conditions. Additionally, mainstream brands have been able to offer amenities and devices such as leather, wood, and anti-lock brakes, previously found only on luxury cars, as the costs decline.

However, luxury vehicle sales remained relatively high compared to their non-luxury counterparts. This was aided by growing interest in luxury vehicles from emerging markets such as China and Russia.

Sales in the entry-level luxury segment remained strong throughout the GFC, due to prices being lowered to compete with well-equipped non-luxury cars. For example, in Canada, several luxury manufacturers set sales records in August 2009, due mostly to discounted pricing on entry-level luxury vehicles.

Brands

Some auto manufacturers market their luxury models using the same marque as the rest of their line. Others have created a separate marque (e.g. Lexus, launched by Toyota in 1989) or purchased one (e.g. Bentley, by Volkswagen in 1998).

Occasionally, a luxury car is initially sold under a mainstream marque and is later rebranded under a specific luxury marque (for example, the Hyundai's Genesis to Genesis G80 as well as the Citroën DS to DS 5).

For mass-produced luxury cars, sharing of platforms or components with other models is common, as per modern automotive industry practice.

Subcompact executive / premium compact
The premium compact class is the category of the smallest luxury cars. It became popular in the mid-2000s, when European manufacturers (such as Audi, Volvo, BMW, and Mercedes-Benz) introduced new entry-level models that were smaller and cheaper than their compact executive models. The premium compact cars are usually based on the platform of a compact car (also known as "small family car" or C-segment), while some models may be based on a subcompact car (also known as supermini or B-segment).

Examples include the Mercedes-Benz A-Class and CLA-Class, Audi A3, Volvo S40, BMW 1 Series, and 2 Series. Premium compacts compete with well-equipped mid-size cars, and highly optioned premium compact cars can have pricing and features that overlap models in the compact executive segment.

Compact executive / compact luxury
A compact executive car or a compact luxury car is a premium car larger than a premium compact and smaller than an executive car. In European classification, compact executive cars are part of the D-segment. In North American terms, close equivalents are "compact premium car", "compact luxury car", "entry-level luxury car" and "near-luxury car". Compact executive cars are usually based on the platform of a mid-size car (also known as large family car or D-segment), while some models may be based on a compact car (also known as small family car or C-segment).

Executive / mid-size luxury
Executive car is a British term for an automobile larger than a large family car. In official use, the term is adopted by Euro NCAP, a European organization founded to test for car safety. It is a passenger car classification defined by the European Commission.

Luxury saloon / full-size luxury sedan
The next category of luxury cars is known in Great Britain as a "luxury saloon" or "luxury limousine,"   and is known in the United States as a full-size luxury sedan, large luxury sedan, or flagship sedan. It is the equivalent of the European F-segment and the German Oberklasse segment.

Many of these luxury saloons are the flagship for the marque and include the newest automotive technology. Several models are available in long-wheelbase versions, which provide additional rear legroom and may have a higher level of standard features.

Examples of luxury saloons / full-size luxury sedans include the BMW 7 Series, Jaguar XJ, Cadillac CT6, Genesis G90, Audi A8, Mercedes-Benz S-Class, Lexus LS, Hongqi H9, Porsche Panamera and Maserati Quattroporte.

Ultra-luxury
Luxury cars costing over US$100,000 (as of 2007) can be considered as "ultra-luxury cars". Examples include the Rolls-Royce Phantom, Maybach 57 and 62, Hongqi L5, Bentley Mulsanne, Toyota Century, and Aurus Senat. High-end sports cars which are targeted towards performance rather than luxury are not usually classified as ultra-luxury cars, even when their cost is greater than US$100000. The history of a brand and the exclusivity of a particular model can result in price premiums compared to luxury cars with similar features from less prestigious manufacturers. Ultra-luxury cars are usually selected as the official state car.

Grand tourer
Grand tourers are essentially high-performance vehicles. These vehicles are generally two-door coupes and are made for long-distance driving, combined with the luxury of an executive car or full-size luxury car.

Luxury SUV / crossover
Long before the luxury SUV segment was defined and became popular, the 1966 Jeep Super Wagoneer was marketed at the time as a fully-equipped station wagon. It was the first off-road SUV to offer a V8 engine and automatic transmission along with luxury car trim and equipment. Standard features included bucket seating, a center console, air conditioning, a seven-position tilt steering wheel, a vinyl roof, and gold-colored trim panels on the body sides and tailgate. By the late 1970s, optional equipment included an electric sunroof. The 1978 Jeep Wagoneer Limited was the spiritual successor to the Super Wagoneer and was the first four-wheel drive car to use leather upholstery.

The Range Rover was released in 1970 as a two-door vehicle for off-road durability with few "creature comforts." A four-door version was added in 1981, and the model was pushed upmarket in 1983 by introducing an automatic transmission (Chrysler's A727 TorqueFlite) as an option. The Range Rover had a long-travel coil-spring suspension and an aluminium V8 engine.

In the mid-1990s, the SUV market expanded with new entrants. By the mid-1990s, the entry-level Ford Explorer and upscale Jeep Grand Cherokee were the market leaders for SUVs. The fastest-growing sector of this market was for the so-called luxury SUVs, which included the Jeep Grand Cherokee ... the Grand Cherokee's allure: "This vehicle is proof you can have a true off-road vehicle without giving up luxuries and amenities" with the Jeep providing a crucial new intangible factor for buyers—image.

The SUV models generated higher profit margins than passenger cars, and car manufacturers began introducing new luxury SUVs during the late 1990s. SUVs such as the 1995 Lexus LX, 1997 Mercedes-Benz M-Class, and 1998 Lincoln Navigator were the first SUVs produced by these luxury car brands. Some of these early luxury SUV models used unibody construction, becoming part of the trend moving away from the body-on-frame construction traditionally used by off-road vehicles.

During the mid-2000s, SUVs from luxury car brands grew by almost 40% in the United States to more than 430,000 vehicles (excluding SUV-only brands like Hummer and Land Rover), at a time when luxury car sales suffered a 1% decline, and non-luxury SUV sales were flat. By 2004, 30% of major luxury brands' U.S. sales were SUVs. Crossover SUVs became increasingly popular in the mid-2000s, and manufacturers also began to produce luxury versions of crossovers. The Lexus RX was the earliest luxury crossover on the market, and it has since been the best-selling luxury vehicle in the US. Some luxury crossovers are built on a platform shared with sedans or hatchbacks. For example, the Infiniti FX is based upon the same platform as the Infiniti G35 sedans and coupes. While early luxury crossovers released in the late 1990s have resembled traditional boxy SUVs, later crossovers, such as the Infiniti FX and BMW X6, have been designed with a sporting appearance.

Despite the increased popularity of crossover models, traditional luxury SUVs remain in production. Examples include the Lexus LX, Infiniti QX80, and Lincoln Navigator.

Research data from the mid-2000s suggested that luxury SUV buyers did not consider traditional luxury cars (e.g. sedans and coupes), therefore the SUV is becoming the key to bringing new customers into luxury dealerships.

Luxury car companies have increasingly introduced SUV or crossover models in the 2010s. For example, Rolls-Royce Cullinan, Bentley Bentayga, Aston Martin DBX, Maserati Levante, and Lamborghini Urus. Ferrari has announced that they will release an SUV model Ferrari Purosangue in 2023. Some brands, such as Lincoln, have even moved to an all SUV and/or crossover lineup.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Luxury car


text:
Limo

Limo may refer to:

 * Limousine, a luxury car
 * Limo (woreda), an administrative division in southern Ethiopia
 * Limo, Depok, a district of Depok, West Java, Indonesia
 * Limo, Les Anglais, Haiti, a village in the Les Anglais commune of Haiti
 * LiMo Platform, a mobile phone and device operating system
 * LiMo Foundation, the organization that develops the LiMo Platform
 * Limo, Ghana, a community in Kumbungu District in the Northern Region of Ghana

People
 * Limo (name), a Kalenjin name common in Kenya
 * Benjamin Limo (born 1974), Kenyan long-distance runner and 2005 World Champion over 5000 metres
 * Felix Limo (born 1980), Kenyan long-distance runner and Chicago and London marathon winner
 * Philemon Limo (born 1985), Kenyan long-distance runner competing in 10K races
 * Remmy Limo (born 1971), Kenyan triple jumper
 * Richard Limo (born 1980, a Kenyan long-distance runner and 2001 World Champion over 5000 metres

title: Limo


text:
Talk:Luxury car

VOLVO
An another point that shows that Volvo is a luxury brand even when we talk about the US market is the fact that its dealerships are in the 6th place among the luxury brands...

European size segment
Consider adding in the first paragraph, that these cars are also known as the F-segment across Europe.

Small Image Update
Hi all, I just made a few minor modifications to the page. These include: 1. I added more info about the run-off between the Mark Series and Eldorado in the history section. This is specifically because of the prominence in American culture. 2. I added a photo of Arnold Palmer standing in front of a Mark III and removed the old Chrysler imperial photo to highlight the importance of personal luxury cars in the era, both with the general population and celebrities. 3. I removed the Nash Ambassador photo. The reason for this is because the "brands" section featured over-representation from American models, and unfortunately the Nash is the lesser important of the two. The section appeared cluttered, so it was removed 4. I removed the Hongqi photo. Since the east-asia section is quite small and two photos felt cluttered, the Hongqi was removed and the Toyota Century remained. This is because of the greater international importance of the Toyota.

Please share your thoughts on this as we work to improve the page. Did I make the wrong move? Should we re-add those photos? GeorgeRoush5 (talk) 15:50, 25 August 2022 (UTC)

Missing Luxury Brands
Under Brands I think a few are missing.

Koenigsegg, SSC, W Motors, Tesla, Pagani, Gordon Murray Automotive, Aspark, Hennessey, Delage, Rimac, SP Automotive, Czinger, Deus, Zenvo, Rezvani

Certainly many of these don't have much of a reputation yet and/or have just produced one model, but Pagani, Koenigsegg, and Tesla are very well known and are missing. Haik12 (talk) 01:13, 12 July 2023 (UTC)

 * The majority of what you listed are top end sports car brands - ie they focus on speed and handling. Luxury cars focus on comfort. Price does not equate to luxury.  Stepho  talk 21:04, 12 July 2023 (UTC)
 * The same can be said of many cars on the grand list then, no?
 * If Bugatti is there why aren’t Pagani and Tesla? 50.159.63.75 (talk) 00:24, 13 July 2023 (UTC)

Is this a joke?
The "Classification standards" section has three bullet points that actually could be taken as "classification standards". The bits about "In, these cars are called " are not "classification standards", but mere trivia.

Also, luxury subcompacts? Are you kidding me/us? You even mention "C-segment" when clearly luxury cars are "F-segment". Nobody would consider a Mercedes A-class a luxury car. Yes, you (whoever wrote or rather hacked together this piece of excrement) tried to CYA with "The term is relative and partially subjective, reflecting both objective qualities of a car, as well as the projected and perceived image of the car marque." but that's still just hogwash. Oh, and the reference? This "Economics Web Institute" doesn't even have a Wikipedia article, for crying out loud! A marque does not a luxury car make. jae (talk) 02:03, 22 October 2023 (UTC)

title: Talk:Luxury car


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Agricultural Research Service

The Agricultural Research Service (ARS) is the principal in-house research agency of the United States Department of Agriculture (USDA). ARS is one of four agencies in USDA's Research, Education and Economics mission area. ARS is charged with extending the nation's scientific knowledge and solving agricultural problems through its four national program areas: nutrition, food safety and quality; animal production and protection; natural resources and sustainable agricultural systems; and crop production and protection. ARS research focuses on solving problems affecting Americans every day. The ARS Headquarters is located in the Jamie L. Whitten Building on Independence Avenue in Washington, D.C., and the headquarters staff is located at the George Washington Carver Center (GWCC) in Beltsville, Maryland. For 2018, its budget was $1.2 billion.

Mission
ARS conducts scientific research for the American public. Their main focus is on research to develop solutions to agricultural problems and provide information access and dissemination to:
 * ensure high quality, safe food and other agricultural products,
 * assess the nutritional needs of Americans,
 * sustain a competitive agricultural economy.
 * enhance the natural resource base and the environment, and
 * provide economic opportunities to rural citizens, communities, and society as a whole.

ARS research complements the work of state colleges and universities, agricultural experiment stations, other federal and state agencies, and the private sector. ARS research may often focus on regional issues that have national implications, and where there is a clear federal role. ARS also provides information on its research results to USDA action and regulatory agencies and to several other federal regulatory agencies, including the Food and Drug Administration and the United States Environmental Protection Agency.

ARS disseminates much of its research results through scientific journals, technical publications, Agricultural Research magazine, and other forums. Information is also distributed through ARS's National Agricultural Library (NAL). ARS has more than 150 librarians and other information specialists who work at two NAL locations—the Abraham Lincoln Building in Beltsville, Maryland; and the DC Reference Center in Washington, D.C. NAL provides reference and information services, document delivery, interlibrary loan and interlibrary borrowing services to a variety of audiences.

History
Prior to the inception of ARS, agricultural research was first conducted under the umbrella of the Agricultural Department in the U.S. Patent Office in 1839. It was created to collect statistics, distribute seeds and compile and distribute pertinent information. In 1862 the USDA was created and agricultural research was moved to its department. That same year, the department issued its first research bulletin on the sugar content of grape varietals and their suitability for wine. Six years later the USDA would begin its first research on animal diseases, specifically hog cholera, which was causing devastating losses at the time. In the early 1900s the USDA began analyzing food composition and the first studies of nutrition and the effects of cooking and processing foods were conducted. Finally, in 1953 the Agricultural Research Service was created to be the USDA's primary scientific research agency.

Research centers
ARS supports more than 2,000 scientists and post docs working on approximately 690 research projects within 15 National Programs at more than 90 research locations. The ARS is divided into 5 geographic areas: Midwest Area, Northeast Area, Pacific West Area, Plains Area, and Southeast Area. ARS has five major regional research centers:
 * Western Regional Research Center in Albany, California
 * Center for Agricultural Resources Research in Fort Collins, Colorado
 * Southern Regional Research Center in New Orleans, Louisiana
 * National Center for Agricultural Utilization Research in Peoria, Illinois
 * Eastern Regional Research Center in Wyndmoor, Pennsylvania

The research centers focus on innovation in agricultural practices, pest control, health, and nutrition among other things. Work at these facilities has given life to numerous products, processes, and technologies.

ARS' Henry A. Wallace Beltsville Agricultural Research Center (BARC) in Beltsville, Maryland, is the world's largest agricultural research complex. Other D.C. area locations include the United States National Agricultural Library and the United States National Arboretum.

ARS also has six major human nutrition research centers that focus on solving a wide spectrum of human nutrition questions by providing authoritative, peer-reviewed, science-based evidence. The centers are located in Arkansas, Maryland, Texas, North Dakota, Massachusetts (the Human Nutrition Research Center on Aging), and California. ARS scientists at these centers study the role of food and dietary components in human health from conception to advanced age.

The ARS also offers the Culture Collection, which is the largest public collection of microorganisms in the world, containing approximately 93,000 strains of bacteria and fungi. The ARS Culture Collection is housed at the National Center for Agricultural Utilization Research (NCAUR) ARS operates the U.S. Horticultural Research Laboratory in Fort Pierce, Florida, and the U.S. National Poultry Research Center in Athens, Georgia. Other notable ARS facilities include Northern Great Basin Experimental Range in Oregon, and formerly the Plum Island Animal Disease Center off Long Island.

Several ARS research units focus on pests, diseases, and management practices of horticultural crops. The is located on the campus of the University of Minnesota in Saint Paul. It primarily hosts research into rusts and Fusaria of cereals. The San Joaquin Valley Agricultural Sciences Center in Parlier, California conducts research on specialty crops including grapes, citrus, almonds, alfalfa, peaches, pomegranates, and many others. Several research locations throughout the US also house germplasm collections of plant species important for agricultural and industrial uses as part of the National Genetic Resources Program.

Research impacts
From the very beginning the Department of Agriculture and in turn the Agricultural Research Service has been focused on improving not only the farming industry but also the quality of food and the health of Americans. In 1985, technology to produce lactose-free milk, yogurt, and ice cream was developed through the Agricultural Research Service. The grape breeding program, currently located in Parlier, CA which began in 1923, developed seedless grapes and continues to release new grapevine varieties with improved traits. The ARS Citrus and Subtropical Products Laboratory in Winter Haven, Florida, actively works to improve the taste of orange juice concentrate.

The Agricultural Research Service had a Toxoplasma gondii (T. gondii) research program, which experimented on cats infected with the parasite, from 1982 until 2018. The prevalence of T. gondii parasite has been reduced by 50% in the U.S. which the ARS claims is a result of their research. The USDA has since discontinued the use of cats in their research amid acute accusations of animal abuse, and a lack of meaningful contribution to the field in recent years (only three of thirteen papers published about T. gondii by the ARS were published after the year 2000).

More recently, the ARS has focused research on genetics and plant and animal DNA. Their research has developed pest-resistant corn, faster growing plants and fish, and a focus on plant and animal genome research and mapping. Outside of scientific research, the ARS has worked to release databases on food components in order to assist consumers with making informed decisions about food choices.

title: Agricultural Research Service


text:
Alanine transaminase

Alanine transaminase (ALT) is a transaminase enzyme. It is also called alanine aminotransferase (ALT or ALAT) and was formerly called serum glutamate-pyruvate transaminase or serum glutamic-pyruvic transaminase (SGPT) and was first characterized in the mid-1950s by Arthur Karmen and colleagues. ALT is found in plasma and in various body tissues but is most common in the liver. It catalyzes the two parts of the alanine cycle. Serum ALT level, serum AST (aspartate transaminase) level, and their ratio (AST/ALT ratio) are routinely measured clinically as biomarkers for liver health.

The half-life of ALT in the circulation approximates 47 hours. Aminotransferase is cleared by sinusoidal cells in the liver.

Function
ALT catalyzes the transfer of an amino group from L-alanine to α-ketoglutarate, the products of this reversible transamination reaction being pyruvate and L-glutamate.

 * L-alanine + α-ketoglutarate ⇌ pyruvate + L-glutamate

ALT (and all aminotransferases) require the coenzyme pyridoxal phosphate, which is converted into pyridoxamine in the first phase of the reaction, when an amino acid is converted into a keto acid.

Clinical significance
ALT is commonly measured clinically as part of liver function tests and is a component of the AST/ALT ratio. When used in diagnostics, it is almost always measured in international units/liter (IU/L) or μkat. While sources vary on specific reference range values for patients, 0-40 IU/L is the standard reference range for experimental studies.

Elevated levels
Test results should always be interpreted using the reference range from the laboratory that produced the result. However typical reference intervals for ALT are:

Significantly elevated levels of ALT (SGPT) often suggest the existence of other medical problems such as viral hepatitis, diabetes, congestive heart failure, liver damage, bile duct problems, infectious mononucleosis, or myopathy, so ALT is commonly used as a way of screening for liver problems. Elevated ALT may also be caused by dietary choline deficiency. However, elevated levels of ALT do not automatically mean that medical problems exist. Fluctuation of ALT levels is normal over the course of the day, and they can also increase in response to strenuous physical exercise.

When elevated ALT levels are found in the blood, the possible underlying causes can be further narrowed down by measuring other enzymes. For example, elevated ALT levels due to hepatocyte damage can be distinguished from bile duct problems by measuring alkaline phosphatase. Also, myopathy-related elevations in ALT should be suspected when the aspartate transaminase (AST) is greater than ALT; the possibility of muscle disease causing elevations in liver tests can be further explored by measuring muscle enzymes, including creatine kinase. Many drugs may elevate ALT levels, including zileuton, omega-3 acid ethyl esters (Lovaza), anti-inflammatory drugs, antibiotics, cholesterol medications, some antipsychotics such as risperidone, and anticonvulsants. Paracetamol (acetaminophen) may also elevate ALT levels.

For years, the American Red Cross used ALT testing as part of the battery of tests to ensure the safety of its blood supply by deferring donors with elevated ALT levels. The intent was to identify donors potentially infected with hepatitis C because no specific test for that disease was available at the time. Prior to July 1992, widespread blood donation testing in the US for hepatitis C was not carried out by major blood banks. With the introduction of second-generation ELISA antibody tests for hepatitis C, the Red Cross changed the ALT policy. , donors previously disqualified for elevated ALT levels and no other reason may be reinstated as donors when they contact the donor-counseling department of their regional Red Cross organization.

In 2000, the American Association for Clinical Chemistry determined that the appropriate terminology for AST and ALT are aspartate aminotransferase and alanine aminotransferase. The term transaminase is outdated and no longer used in liver disease.

Low ALT
Low plasma ALT can be a marker of low muscle mass and is associated with frailty, sarcopenia, disability, as well as increased mortality in the elderly population. In patients with inflammatory bowel disease, low ALT is associated with a more active disease.

title: Alanine transaminase


text:
Burnside's lemma

Burnside's lemma, sometimes also called Burnside's counting theorem, the Cauchy–Frobenius lemma, or the orbit-counting theorem, is a result in group theory that is often useful in taking account of symmetry when counting mathematical objects. It was discovered by Augustin Louis Cauchy and Ferdinand Georg Frobenius, and became well-known after William Burnside quoted it. The result enumerates orbits of a symmetry group acting on some objects: that is, it counts distinct objects, considering objects symmetric to each other as the same; or counting distinct objects up to a symmetry equivalence relation; or counting only objects in canonical form. For example, in describing possible organic compounds of certain type, one considers them up to spatial rotation symmetry: different rotated drawings of a given molecule are chemically identical. (However a mirror reflection might give a different compound.)

Formally, let G be a finite group that acts on a set X. For each g in G, let Xg denote the set of elements in X that are fixed by g (left invariant by g): that is, Xg = { x ∈ X | g.x = x }. Burnside's lemma asserts the following formula for the number of orbits, denoted |X/G|:

$$|X/G| = \frac{1}{|G|}\sum_{g \in G}|X^g|.$$

Thus the number of orbits (a natural number or +∞) is equal to the average number of points fixed by an element of G. For an infinite group G, there is still a bijection:

$$G \times X/G \ \longleftrightarrow\ \coprod_{g \in G} X^g. $$

Necklaces
There are 8 possible bit strings of length 3, but tying together the string ends gives only four distinct 2-colored necklaces of length 3, given by the canonical forms 000, 001, 011, 111: the other strings 100 and 010 are equivalent to 001 by rotation, while 110 and 101 are equivalent to 011. That is, rotation equivalence splits the set X of strings into four orbits: "$X = \{000\}\cup \{001, 010, 100\}\cup \{011, 101, 110\}\cup \{111\}.$"The Burnside formula uses the number of rotations, which is 3 including the null rotation, and the number of bit strings left unchanged by each rotation. All 8 bit vectors are unchanged by the null rotation, and two (000 and 111) are unchanged by the other two rotations. Thus the number of orbits is: $$4 = \frac{1}{3}(8+2+2).$$

For length 4, there are 16 possible bit strings; 4 rotations; the null rotation leaves all 16 strings unchanged; the 1-rotation and 3-rotation each leave two strings unchanged (0000 and 1111); the 2-rotation leaves 4 bit strings unchanged (0000, 0101, 1010, 1111). The number of distinct necklaces is thus: $$6 = \tfrac{1}{4}(16+2+4+2)$$, represented by the canonical forms 0000, 0001, 0011, 0101, 0111, 1111.

The general case of n bits and k colors is given by a necklace polynomial.

Colorings of a cube
Burnside's lemma can compute the number of rotationally distinct colourings of the faces of a cube using three colours.

Let X be the set of 36 possible face color combinations that can be applied to a fixed cube, and let the rotation group G of the cube act on X by moving the colored faces: two colorings in X belong to the same orbit precisely when one is a rotation of the other. Rotationally distinct colorings correspond to group orbits, and can be found by counting the sizes of the fixed sets for the 24 elements of G, the colorings left unchanged by each rotation:

 * the identity element fixes all 36 colorings
 * six 90-degree face rotations each fix 33 colorings
 * three 180-degree face rotations each fix 34 colorings
 * eight 120-degree vertex rotations each fix 32 colorings
 * six 180-degree edge rotations each fix 33 colorings.

A detailed examination may be found here.

The average fixed-set size is thus:

 * $$ |X/G| = \frac{1}{24}\left(3^6+6\cdot 3^3 + 3 \cdot 3^4 + 8 \cdot 3^2 + 6 \cdot 3^3 \right) = 57. $$

There are 57 rotationally distinct colourings of the faces of a cube in three colours. In general, the number of rotationally distinct colorings of the faces of a cube in n colors is:

 * $$ \frac{1}{24}\left(n^6+3n^4 + 12n^3 + 8n^2\right). $$

Proof
In the proof of Burnside's lemma, the first step is to re-express the sum over the group elements g ∈ G as an equivalent sum over the set of elements x ∈ X:

 * $$\sum_{g \in G}|X^g| = \#\{(g,x)\in G\times X \mid g \cdot x = x\} = \sum_{x \in X} |G_x|.$$

Here Xg = {x ∈ X | g.x = x} is the set of points of X fixed by g ∈ G, whereas Gx = {g ∈ G | g.x = x} is the stabilizer subgroup of G, symmetries that fix the point x ∈ X.)

The orbit-stabilizer theorem says that for each x ∈ X there is a natural bijection between the orbit G·x = {g·x | g ∈ G} and the set of left cosets G/Gx. Lagrange's theorem implies:

 * $$|G \cdot x| = [G\,:\,G_x] = |G| / |G_x|.$$

The sum may therefore be rewritten as:

 * $$\sum_{x \in X} |G_x| = \sum_{x \in X} \frac{|G|}{|G \cdot x|} = |G| \sum_{x \in X}\frac{1}{|G \cdot x|}.$$

Writing X as the disjoint union of its orbits in X/G:

 * $$|G|\, \sum_{x \in X}\frac{1}{|G \cdot x|}

\ =\ |G|\!\sum_{A\in X/G}\, \sum_{x\in A} \frac{1}{|A|} \ =\ |G|\! \sum_{A\in X/G} 1 \ =\ |G|\,|X/G|. $$

Putting everything together gives the desired result:

 * $$\sum_{g \in G}|X^g| = |G| \cdot |X/G|.$$

This is similar to the proof of the conjugacy class equation, which considers the conjugation action of G on itself: X = G and g.x = gxg−1, so that the stabilizer of x is centralizer: Gx = ZG(x).

Enumeration vs. generation
Burnside's lemma counts distinct objects, but it does not construct them. In general, combinatorial generation with isomorph rejection considers the symmetries of g, on objects x. But instead of checking that g.x = x, it checks that g.x has not already been generated. One way to accomplish this is by checking that g.x is not lexicographically less than x, using the lexicographically least member of each equivalence class as the canonical form of the class. Counting the objects generated with such a technique can verify that Burnside's lemma was correctly applied.

History: the lemma that is not Burnside's
William Burnside stated and proved this lemma in his 1897 book on finite groups, attributing it to. But even prior to Frobenius, the formula was known to Cauchy in 1845. Consequently, this lemma is sometimes referred to as the lemma that is not Burnside's. Misnaming scientific discoveries is referred to as Stigler's law of eponymy.

title: Burnside's lemma