Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates,[1][2] and indeed a premier example of a complex evolutionary novelty.[3] They are among the characteristics that distinguish the extant Aves from other living groups. Feathers have also been noticed in those Theropoda which have been termed feathered dinosaurs. Although feathers cover most parts of the body of birds, they arise only from certain well-defined tracts on the skin. They aid in flight, thermal insulation, waterproofing and coloration that helps in communication and protection.[4]
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Feathers are among the most complex integumentary appendages found in vertebrates and are formed in tiny follicles in the epidermis, or outer skin layer, that produce keratin proteins. The β-keratins in feathers, beaks and claws — and the claws, scales and shells of reptiles — are composed of protein strands hydrogen-bonded into β-pleated sheets, which are then further twisted and crosslinked by disulfide bridges into structures even tougher than the α-keratins of mammalian hair, horns and hoof.[5][6] The exact signals that induce the growth of feathers on the skin are not known but it has been found that the transcription factor cDermo-1 induces the growth of feathers on skin and scales on the leg.[7]
There are two basic types of feather: vaned feathers which cover the exterior of the body, and down feathers which are underneath the vaned feathers. The pennaceous feathers are vaned feathers. Also called contour feathers, pennaceous feathers arise from tracts and cover the whole body. A third rarer type of feathers, filoplumes, is hairlike and (if present in a bird) grows along the fluffy down feathers. In some passerines, filoplumes arise exposed beyond the contour feathers on the neck.[1] The remiges, or flight feathers of the wing, and rectrices, the flight feathers of the tail are the most important feathers for flight. A typical vaned feather features a main shaft, called the rachis. Fused to the rachis are a series of branches, or barbs; the barbs themselves are also branched and form the barbules. These barbules have minute hooks called barbicels for cross-attachment. Down feathers are fluffy because they lack barbicels, so the barbules float free of each other, allowing the down to trap much air and provide excellent thermal insulation. At the base of the feather, the rachis expands to form the hollow tubular calamus (or quill) which inserts into a follicle in the skin. The basal part of the calamus is without vanes. This part is embedded within the skin follicle and has an opening at the base (proximal umbilicus) and a small opening on the side (distal umbilicus).[8]
Hatchling birds of some species have a special kind of natal down (neossoptiles) and these are pushed out when the normal feathers (teleoptiles) emerge.[1]
Flight feathers are stiffened so as to work against the air in the downstroke but yield in other directions. It is noted that the pattern of orientation of β-keratin fibers in the feathers of flying birds differs from that in flightless birds. The fibers are better aligned in the middle of the feather and less aligned towards the tips.[9][10]
Feathers insulate birds from water and cold temperatures. They may also be plucked to line the nest and provide insulation to the eggs and young. The individual feathers in the wings and tail play important roles in controlling flight. Some species have a crest of feathers on their heads. Although feathers are light, a bird's plumage weighs two or three times more than its skeleton, since many bones are hollow and contain air sacs. Color patterns serve as camouflage against predators for birds in their habitats, and by predators looking for a meal. As with fish, the top and bottom colors may be different to provide camouflage during flight. Striking differences in feather patterns and colors are part of the sexual dimorphism of many bird species and are particularly important in selection of mating pairs. In some cases there are differences in the UV reflectivity of feathers across sexes even though no differences in color are noted in the visible range.[11] The wing feathers of male Club-winged Manakins Machaeropterus deliciosus have special structures that are used to produce sounds by stridulation.[12]
Some birds have a supply of powder down feathers which grow continuously, with small particles regularly breaking off from the ends of the barbules. These particles produce a powder that sifts through the feathers on the bird's body and acts as a waterproofing agent and a feather conditioner. Powder down has evolved independently in several taxa and can be found in down as well as pennaceous feathers. They may be scattered in plumage in the pigeons and parrots or in localized patches on the breast, belly or flanks as in herons and frogmouths. Herons use their bill to break the feathers and to spread them while cockatoos may use their head as a powder puff to apply the powder.[13] Waterproofing can be lost by exposure to emulsifying agents due to human pollution. Feathers can become waterlogged and birds may sink. It is also very difficult to clean and rescue birds whose feathers have been fouled by oil spills. The feathers of cormorants soak up water and help in reducing buoyancy and thereby allowing the birds to swim submerged.[14]
Bristles are stiff, tapering feathers with a large rachis but few barbs. Rictal bristles are bristles found around the eyes and bill. They may serve a similar purpose to eyelashes and vibrissae in mammals. It has been suggested that they may aid insectivorous birds in prey capture or that it may have sensory functions, however there is no clear evidence.[15] In one study, Willow Flycatchers (Empidonax traillii) were found to catch insects equally well before and after removal of the rictal bristles.[16]
Grebes are peculiar in their habit of ingesting their own feathers and also feeding them to their young. Observations on the diet and feather eating frequency suggest that ingesting feathers particularly down from their flanks aids in forming easily ejectable pellets along with their diet of fish.[17]
Contour feathers are not uniformly distributed on the skin of the bird except in some groups such as the Penguins, ratites and screamers.[18] In most birds the feathers grow from specific tracts of skin called pterylae while there are regions which are free of feathers called apterylae. Filoplumes and down may arise from the apteriae, regions between the pterylae. The arrangement of these feather tracts, pterylosis or pterylography, varies across bird families and has been used in the past as a means for determining the evolutionary relationships of bird families.[19][20]
The colors of feathers are produced by the presence of pigments, or by microscopic refractive structures, or by a combination of both.
Most feather pigments are melanins (brown and beige pheomelanins, black and grey eumelanins) and carotenoids (red, yellow, orange); other pigments occur only in certain taxa – the yellow to red psittacofulvins[21] (found in some parrots) and the red turacin and green turacoverdin (porphyrin pigments found only in turacos). Structural coloration[4][22][23] is involved in the production of blue colors, iridescence, most ultraviolet reflectance and in the enhancement of pigmentary colors; structural iridescence has been reported[24] in fossil feathers dating back 40 million years. White feathers lack pigment and scatter light diffusely; albinism in birds is caused by defective pigment production, though structural coloration will not be affected (as can be seen e.g. in blue-and-white budgerigars).
For example, the blues and bright greens of many parrots are produced by constructive interference of light reflecting from different layers of the structures in feathers, in the case of green plumage in addition to the yellow pigments; the specific feather structure involved is sometimes called the Dyck texture.[25][26] Melanin is often involved in the absorption of some of the light; in combination with yellow pigment it produces dull olive-greens.
In some birds, the feather colors may be created or altered by uropygial gland secretions. The yellow bill colors of many hornbills are produced by preen gland secretions. Other differences that may only be visible in the ultraviolet region have been suggested[13] but studies have failed to find evidence.[27] Uropygial oil secretion may also have an inhibitory effect on feather bacteria.[28]
A bird's feathers undergo wear and tear and are replaced periodically during its life through molting. New feathers, known as blood, or pin feathers (depending on the stage of growth) when developing, are formed through the same follicle from which the old ones were fledged. The presence of melanin in feathers increases their resistance to abrasion.[29] One study notes that melanin based feathers were observed to degrade more quickly under bacterial action, even compared to unpigmented feathers from the same species, than those unpigmented or with carotenoid pigments.[30] However, another study the same year compared the action of bacteria on pigmentations of two song sparrow species and observed that the darker pigmented feathers were more resistant and they cited other research also published in 2004 that stated increased melanin provided greater resistance. They observed that the greater resistance of the darker birds confirmed Gloger's rule.[31] The evolution of coloration is based on sexual selection and it has been suggested that carotenoid-based pigments may have evolved since they are likely to be more honest signals of fitness because they are derived from special diets,[32][33] or because carotenoids are also required for immune function.[34]
The feather surface is the home for some ectoparasites, notably feather lice (Phthiraptera) and feather mites. Feather lice typically live on a single host and can move only from parents to chicks or mating birds and occasionally by phoresy. This life history has resulted in most of the species being specific to the host and coevolving with the host, making them of interest in phylogenetic studies.[35]
Feather holes are chewing traces of lice (most probably Brueelia spp. lice) on the wing and tail feathers. They were described on barn swallows, and because of easy countability, many evolutionary, ecological, and behavioral publications use them to quantify the intensity of infestation.
Interestingly, parasitic cuckoos which grow up in the nests of other species also have host specific feather lice and these seem to be transmitted only after they leave the host nest.[36]
Birds maintain their feather condition by bathing in water, dust bathing and preening. A peculiar behavior of birds, anting, where ants are introduced into the plumage was suggested to help in reducing parasites but no supporting evidence has been found.[37]
Feathers have a number of utilitarian, cultural and religious uses.
Feathers are both soft and excellent at trapping heat; thus, they are sometimes used in high-class bedding, especially pillows, blankets, and mattresses. They are also used as filling for winter clothing, such as quilted coats and sleeping bags; goose and eider down have great loft, the ability to expand from a compressed, stored state to trap large amounts of compartmentalized, insulating air.[38]
Bird feathers have long been used for fletching arrows. Colorful feathers such as those belonging to pheasants have been used to decorate fishing lures.
Feathers of large birds (most often geese) have been and are used to make quill pens. The word pen itself is derived from the Latin penna for feather.[39] In French, plume can mean either feather or pen.
Feathers are also valuable in aiding the identification of species in forensic studies, particularly in bird strikes to aircraft. The ratios of hydrogen isotopes in feathers help in determining the geographic origins of birds.[40] Feathers may also be useful in the non-destructive sampling of pollutants.[41]
The poultry industry produces a large amount of feathers as waste, and like other forms of keratin, these are slow in their decomposition. Feather waste has been used in a number of industrial applications as a medium for culturing microbes,[42] biodegradeable polymers,[43] and production of enzymes.[44] Feather proteins have been tried as an adhesive for wood board.[45]
Eagle feathers have great cultural and spiritual value to American Indians in the USA and First Nations peoples in Canada as religious objects. In the United States the religious use of eagle and hawk feathers are governed by the eagle feather law, a federal law limiting the possession of eagle feathers to certified and enrolled members of federally recognized Native American tribes.
Various birds and their plumages serve as cultural icons throughout the world, from the hawk in ancient Egypt to the bald eagle and the turkey (bird) in the United States. In Greek mythology, Daedelus the inventor and Icarus tried to escape his prison by attaching feathered wings to his shoulders with wax, which was melted by the Sun.
In South America, brews made from the feathers of Condors are used in traditional medications.[46] In India, feathers of the Indian Peacock have been used in traditional medicine for snakebite, infertility and coughs.[47][48]
During the 18th, 19th, and even 20th Centuries a booming international trade in plumes, to satisfy market demand in North America and Europe for extravagant head-dresses as adornment for fashionable women, caused so much destruction (for example, to egret breeding colonies) that a major campaign against it by conservationists led to the Lacey Act and caused the fashion to change and the market to finally collapse. Frank Chapman noted in 1886 that as many as 40 species of birds were used in about three-fourths of the 700 ladies' hats that he observed in New York City.[49][50][51]
More recently, rooster plumage has become a popular trend as a hairstyle accessory, with feathers formerly used solely as fishing lures being now used to provide color and style to hair.[52]
The functional view on the evolution of feathers has traditionally focused on insulation, flight and display. Discoveries of non-flying Late Cretaceous feathered dinosaurs in China however suggest that flight could not have been the original primary function.[53] There have been suggestions that feathers may have had their original function in thermoregulation, waterproofing or even as sinks for metabolic wastes such as sulphur.[54] While feathers have been suggested as having evolved from reptilian scales, there are numerous objections, and more recent explanations have arisen from the paradigm of evolutionary developmental biology.[2] Theories of the scale-based origins of feathers suggest that the planar scale structure was modified for their development into feathers by splitting to form the webbing; however, the developmental process involves a tubular structure arising from a follicle and the tube splitting longitudinally to form the webbing.[1][2] The number of feathers per unit area of skin is higher in smaller birds than in larger birds, and this trend indicates their important role in thermal insulation, since smaller birds lose more heat due to the relatively larger surface area in proportion to their body weight.[4] The miniaturization of birds also played a role in the evolution of powered flight.[55] The coloration of feathers is believed to be primarily evolved in response to sexual selection. In many cases the physiological condition of the birds (especially males) is indicated by the quality of their feathers and this is used (by the females) in mate choice.[56][57]
Several non-avian dinosaurs had feathers on their limbs that would not have functioned for flight.[2] One theory is that feathers originally evolved on dinosaurs as a result of insulation properties; those small dinosaurs that then grew longer feathers may have found them helpful in gliding, leading to the evolution of proto-birds like Archaeopteryx and Microraptor zhaoianus. Dinosaurs that had feathers or protofeathers include Pedopenna daohugouensis, [58] and Dilong paradoxus, a tyrannosauroid which is 60 to 70 million years older than Tyrannosaurus rex.[59]
The majority of dinosaurs known to have had feathers or protofeathers are saurischians, however featherlike "filamentous integumentary structures" are also known from the ornithischians Tianyulong and Psittacosaurus.[60] The exact nature of these structures is still under study. However, it is believed that the stage 1 feathers such as those seen in these two ornithischians likely functioned in display. [61]
Since the 1990s, dozens of feathered dinosaurs have been discovered in the clade Maniraptora, which includes the clade Avialae and the recent common ancestors of birds, Oviraptorosauria and Deinonychosauria. In 1998, the discovery of a feathered oviraptorosaurian, Caudipteryx zoui, challenged the notion that feathers were an exclusive structure of Avialae.[62] Buried in the Yixian Formation in Liaoning, China, C. zoui lived during the Early Cretaceous Period. Present on the forelimbs and tails, their integumentary structure has been accepted as pennaceous vaned feathers based on the rachis and herringbone pattern of the barbs. In the clade Deinonychosauria, the continued divergence of feathers is also apparent in the families Troodontidae and Dromaeosauridae. Branched feathers with ranchis, barbs, and barbules were discovered in many members including Sinornithosaurus millenii, a dromaeosaurid found in the Yixian formation (124.6 MYA).[63]
Previously, a temporal paradox existed in the evolution of feathers - theropods with highly derived bird-like characteristics occurred at a later time than Archaeopteryx, suggesting that the descendants of birds arose before the ancestor. However, this paradox was resolved in 2009 with the discovery of Anchiornis huxleyi, found in the Late Jurassic Tiaojishan Formation (160 MYA) in western Liaoning.[64][65] By predating Archaeopteryx, Anchornis proves the existence of a modernly feathered theropod ancestor, providing insight into the dinosaur-bird transition. The specimen shows distribution of large pennaceous feathers on the forelimbs and tail, implying that pennaceous feathers spread to the rest of the body at an earlier stage in theropod evolution.[66] The discovery, in 2011, of feathers preserved in amber, within samples dating to 80 mya, suggests the coexistence of theropods and birds, with both theropod and avian feather types commingled in the samples.[67] x
Several studies of feather development in the embryos of modern birds, coupled with the distribution of feather types among various prehistoric bird precursors, have allowed scientists to attempt a reconstruction of the sequence in which feathers first evolved and developed into the types found on modern birds.
Feather evolution was been broken down into the following stages by Xu and Guo in 2009:[61]
However, Foth (2011) showed that some of these purported stages (stages 2 and 5 in particular) are likely simply artifacts of preservation caused by the way fossil feathers are crushed and the feather remains or imprints are preserved. Foth re-interpreted stage 2 feathers as crushed or misidentified feathers of at least stage 3, and stage 5 feathers as crushed stage 6 feathers.[68]
The following simplified diagram of dinosaur relationships follows these results, and shows the likely distribution of plumaceous (downy) and pennaceous (vaned) feathers among dinosaurs and prehistoric birds. The diagram follows one presented by Xu and Guo (2009) modified with the findings of Foth (2011).[61] The numbers accompanying each name refer to the presence of specific feather stages. Note that 's' indicates the known presence of scales on the body.
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