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, and waterproofing. In addition, coloration helps in communication and protection. Plumology (or plumage science) is the name for the science that is associated with the study of feathers.
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.
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 entire body. A third rarer type of feather, the filoplume, is hairlike and (if present in a bird; they are entirely absent in ratites) are closely associated with and are often entirely hidden by them, with one or two filoplumes attached and sprouting from near the same point of the skin as each contour feather, at least on a bird's head, neck and trunk. 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 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).
Flight feathers are stiffened so as to work against the air in the downstroke but yield in other directions. It has been observed that the orientation pattern of β-keratin fibers in the feathers of flying birds differs from that in flightless birds: the fibers are better aligned along the shaft axis direction towards the tip, and the lateral walls of rachis region show structure of crossed fibers.