The beak or bill is an external rostrum structure found mostly in birds. A beak is used for pecking, grasping, and holding (in probing for food, eating, manipulating and carrying objects, killing prey, or fighting), preening, courtship, and feeding young. The terms beak and rostrum are also used to refer to a similar organ in some non-avian theropods, ornithischians, pterosaurs, cetaceans, dicynodonts, rhynchosaurs, anuran tadpoles, turtles, monotremes (i.e. echidnas and platypuses, which have a bill-like structure), sirens, pufferfish, billfishes, parrotfish and cephalopods. Although beaks vary significantly in size, shape, color and texture, they share a similar underlying structure. Two bony projections–the upper and lower mandibles–are covered with a thin keratinized layer of epidermis known as the rhamphotheca. In most species, two holes called nares lead to the respiratory system.
Etymology Although the word "beak" was, in the past, generally restricted to the sharpened bills of birds of prey, in modern ornithology, the terms beak and bill are generally considered to be synonymous. The word, which dates from the 13th century, comes from the Middle English bec (via Anglo-French), which itself comes from the Latin beccus.
Anatomy
Although beaks vary significantly in size and shape from species to species, their underlying structures have a similar pattern. All beaks are composed of two jaws, generally known as the maxilla (upper) and mandible (lower). The upper, and in some cases the lower, mandibles are strengthened internally by a complex three-dimensional network of bony spicules (or trabeculae) seated in soft connective tissue and surrounded by the hard outer layers of the beak. The avian jaw apparatus is made up of two units: one four-bar linkage mechanism and one five-bar linkage mechanism.
Mandibles
The upper mandible is supported by a three-pronged bone called the intermaxillary. The upper prong of this bone is embedded into the forehead, while the two lower prongs attach to the sides of the skull. At the base of the upper mandible a thin sheet of nasal bones is attached to the skull at the nasofrontal hinge, which gives mobility to the upper mandible, allowing it to move upward and downward.
The base of the upper mandible, or the roof when seen from the mouth, is the palate; the palate's structure differs greatly in the ratites. Here, the vomer is large and connects with premaxillae and maxillopalatine bones in a condition termed as a "paleognathous palate". All other extant birds have a narrow forked vomer that does not connect with other bones and is then termed as neognathous. The shape of these bones varies across the bird families. The lower mandible is supported by a bone known as the inferior maxillary bone—a compound bone composed of two distinct ossified pieces. These ossified plates (or rami), which can be U-shaped or V-shaped, join distally (the exact location of the joint depends on the species) but are separated proximally, attaching on either side of the head to the quadrate bone. The jaw muscles, which allow the bird to close its beak, attach to the proximal end of the lower mandible and to the bird's skull. The muscles that depress the lower mandible are usually weak, except in a few birds such as the starlings and the extinct huia, which have well-developed digastric muscles that aid in foraging by prying or gaping actions. In most birds, these muscles are relatively small as compared to the jaw muscles of similarly sized mammals.
Rhamphotheca
The outer surface of the beak consists of a thin sheath of keratin called the rhamphotheca, which can be subdivided into the rhinotheca of the upper mandible and the gnathotheca of the lower mandible. The covering arises from the Malpighian layer of the bird's epidermis, growing from plates at the base of each mandible. There is a vascular layer between the rhamphotheca and the deeper layers of the dermis, which is attached directly to the periosteum of the bones of the beak. The rhamphotheca grows continuously in most birds, and in some species, the color varies seasonally. In some alcids, such as the puffins, parts of the rhamphotheca are shed each year after the breeding season, while some pelicans shed a part of the bill called a "bill horn" that develops in the breeding season. While most extant birds have a single seamless rhamphotheca, species in a few families, including the albatrosses and the emu, have compound rhamphothecae that consist of several pieces separated and defined by softer keratinous grooves. Studies have shown that this was the primitive ancestral state of the rhamphotheca, and that the modern simple rhamphotheca resulted from the gradual loss of the defining grooves through evolution.
Tomia
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