Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds. They are considered the most complex integumentary structures found in vertebrates. They are among the outstanding characteristics that distinguish the extant Aves from other living groups. Feathers have also been noticed in Theropoda which have been termed feathered dinosaurs. However, there are also some scientists who disagree with the interpretation of feathered dinosaurs, pointing out instead that birds and some theropods are only convergently similar. 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.
Structure and characteristics
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. 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.
Classification
See also: Flight featherThere 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. 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).
Hatchling birds of some species have a special kind of natal down (neossoptiles) and these are pushed out when the normal feathers (teleoptiles) emerge.
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.
Functions
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. The wing feathers of male Club-winged Manakins Machaeropterus deliciosus have special structures that are used to produce sounds by stridulation.
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. 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.
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. In one study, Willow Flycatchers ( Empidonax traillii ) and they were found to catch insects equally well before and after removal of the rictal bristles.
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.
Distribution
Contour feathers are not uniformly distributed on the skin of the bird except in some groups such as the Penguins, ratites and screamers. 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.
Coloration
The colors of feathers are produced by the presence of pigments such as melanins (browns, blacks, greys), carotenoids (reds, yellows, orange), psittacofulvins (unique red pigments found in some parrots) and porphyrins (such as the green turacoverdin of Turacos) or more often by feather structure. Structural coloration is involved in the production of most greens, blues, iridescent colors, ultraviolet reflectance and in the enhancement of pigmentary colors. Structural iridescence in feathers has been reported in fossil feathers dating back 40 million years.
The blues and greens of many parrots are produced by constructive interference of light reflecting from different layers of the structures in feathers in addition to the yellow carotenoid pigments. Melanin is often involved in the absorption of some of the light in these feathers. The specific feather structure involved is sometimes called the Dyck texture.
Albinism is caused by the lack of pigment in some or all of a bird's feathers.
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 but studies have failed to find evidence. Uropygial
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