Professional Documents
Culture Documents
, 40:461–477 (2000)
PETER R. STETTENHEIM 2
168 Croydon Turnpike, Plainfield, New Hampshire 03781-5403
SYNOPSIS. Avian integument is thin, elastic, and loosely attached to the body,
giving birds the freedom of movement needed for flight. Its epidermis is both
keratinized and lipogenic, and the skin as a whole acts as a sebaceous secretory
organ. The skin is covered by feathers over most of the body, but many birds show
colored bare skin or integumentary outgrowths on the head and neck. Heavily
cornified epidermis covers the beak, claws, spurs, and the scales on the legs and
feet. These structures (except the back of the leg and underside of the foot) contain
beta-keratin like that in reptilian scales. Most birds have sebaceous secretory
461
462 PETER R. STETTENHEIM
amino acid composition, molecular size, gether with the feathers, the skin is more
and organization among the various integ- active in thermoregulation in birds than in
umentary derivatives (Brush, 1980a, b; reptiles. Its surface radiates excess body
Brush and Wyld, 1982; Homberger and heat, absorbs solar radiation, and cools the
Brush, 1986). body by cutaneous water loss (Bernstein,
Avian skin lacks sweat glands and se- 1971; Bartholomew, 1972). The feathering
baceous glands, yet the epidermis itself, in (5feather covering) is a much lighter and
a variety of species produces neutral fats more efficient thermal insulation than rep-
and phospholipids (Lucas, 1968, 1980; tilian scales, owing to the air trapped within
Lavker, 1975). It is strongly lipogenic, in it. This system has presumably evolved
general contrast to reptilian epidermis. along with the evolution of homeothermy
Since avian epidermal cells include both li- (Stevens, 1973).
pogenesis and keratinization in their differ-
tary in young birds. In species such as cer- dian Black Ibises (Pseudibis papillosa), and
tain ibises and storks, areas that are feath- certain curassows, megapodes, and cathartid
ered in young birds subsequently become vultures.
bare, losing feather follicles as well as Wattles, the commonest of soft integu-
feathers. The epidermis of these areas mentary outgrowths, are larger protuber-
shows a high degree of lipoid secretion ances or flaps. They are located on the sides
(personal communication, G. K. Menon). of the head or neck, especially at the base
Once developed by adults, the integumen- of the bill and around the eyes. In some
tary structures are usually permanent, but birds they hang beside the mouth or under
may vary in size or color in relation to the the throat. Wattles occur in cassowaries,
birds’ breeding activity. many cracids, megapodes, ptarmigan,
pheasants, guineafowl, turkeys, rails, jaca-
Throat and neck pouches nas, lapwings, alcids, cotingas, starlings,
mentary thickenings located almost directly jections for holding prey (e.g., mergansers,
below the external ear opening. Histologi- falcons, barbets, toucans). Internally, the
cally, they resemble wattles except that the roof of the mouth also has local thicken-
dermis contains far more collagenic tissue ings, e.g., papillae or spines for holding
(Lucas and Stettenheim, 1972). prey in fish-eating birds, and longitudinal or
Integumentary outgrowths and conspic- transverse ridges for holding and cracking
uous areas of bare skin serve in many cases seeds in cockatoos, parrots, finches and
as visual signals for social or sexual behav- buntings (Ziswiler, 1965; Homberger,
ior. They often indicate physical fitness be- 1980). Nares are located within the basal
cause they change or shrivel when a bird is one-third of the upper bill in most birds and
unhealthy. Thus, they may disclose the con- at the tip of the upper bill in kiwis. They
dition of a potential mate or competitor, in- are usually a simple hole or partly shielded,
dicating dominance, age, or vigor, as in cer- but are tubular in procellariiforms. The
location and number in the beak appear to apodes. Conspicuous by its location and
be related to the way this part is used as a movement, the beak commonly presents vi-
tactile exploratory organ in feeding. They sual signals employed in identification and
are especially numerous in the beak bones displays.
of anseriforms and sandpipers, plus the nail
and edges of the palate in anseriforms and CASQUES
parrots (Berkhoudt, 1980; Gottschaldt, Numerous birds have a hard, bare, heavi-
1985). ly cornified projection known as a casque
on top of their head or their bill. In most
Transitory nestling structures cases, the integument overlies a bony ex-
Embryos or nestlings develop various ac- tension of the skull, but in cassowaries it
cessories on the rhamphotheca that are lost covers a core of tough, elastic, foam-like,
before fledging. In nearly all birds, an egg collagen above the bone. Where the casque
ulate scales). Tarsal scales, not needing These accessories are grown in the fall and
flexibility, are often fused, especially on the molted in spring.
anterior surface.
The two types of scales differ embryo- CLAWS
logically and histochemically as well as Avian claws, like those of reptiles, are
morphologically (Brush and Wyld, 1980). coverings of heavily cornified integument
Scutellate scales contain beta-keratin, the over the bone of a terminal phalanx. All
same as the rhamphotheca and the claws birds have toe claws and many birds have
(Stewart, 1977), whereas reticulate scales wing claws as well. A toe claw is composed
are composed of alpha-keratin. These dif- of a dorsal plate that curves downward on
ferences indicate that they have different the tip and sides and a ventral plate that fills
evolutionary histories. Scutellate scales, be- the space between the sides underneath
ing very similar to crocodilian scales, ap- (Lucas and Stettenheim, 1972). The dorsal
on the feathers (Wrench et al., 1980), their phy and undergo histochemical changes
relative amounts varying among species. which suggest that the secretion serves in
The functions of the uropygial secretion, reproduction, possibly in the mechanics of
although much studied, ought to be re-ex- internal fertilization (Menon et al., 1987).
amined with regard to the skin secretion,
which was long unknown. Together, the two FEATHERS
substances keep the keratin of the integu- Structure
ment and its derivatives flexible and in
good condition. This is particularly impor- Feathers are the most conspicuous, com-
tant for the feathers, whose waterproofing plex, and varied integumentary derivatives
depends chiefly on tiny air bubbles held of birds. They are one of the defining char-
within the meshwork of their barbs (Rijke, acters of the Class Aves, even though they
1970, 1989; Kostina et al., 1996). In order were also present in the early Cretaceous
tangled and matted. In this way, the barbs resist wear and damage because they are
remain fluffy, trapping air in the plumage replaced only infrequently, at regular molts,
for thermal insulation. Additional structural and lost or broken feathers impair flight or
factors influence the fuzziness of the barbs other functions (Bonser, 1996b). In addi-
and hence their insulative value (Lucas and tion, feather parts must have the right de-
Stettenheim, 1972). Plumulaceous barbules gree of stiffness or flexibility according to
are not primitive or deficient as they are their roles. These properties depend on a
sometimes thought to be, but specialized for part’s cross-sectional shape and internal
maintaining downy texture. construction (Rutschke, 1966a) and on the
Pennaceous barbs, though homologous, keratin that constitutes it.
are thicker and stiffer, close together and Feathers are about 90% protein, mostly
parallel, creating vanes that are firm, flat, beta-keratin. This is stronger, yet more flex-
and closely-knit. Their barbules, likewise ible than other avian keratins, owing to dif-
proofing on the ordinary feathers. Powder occur almost exclusively in contour feath-
feathers occur in the Kagu, mesites (Mesi- ers. In addition to their prominent visual
tornithidae), herons and bitterns, pigeons, roles, they are involved with vitamin A in
most cockatoos, and some parrots and sand- vision and possibly with reproduction in the
pipers. yolk. Carotenoids and melanins combine to
Filoplumes are hairlike feathers that do produce colors such as dull green.
not grade into the other types. They have a Porphyrins are iron- or copper-containing
slender, glistening, unpigmented rachis with pigments derived by catabolism in the liver
a few short barbs or barbules at the tip. Fil- and by synthesis from glycine (Brush,
oplumes are always situated close beside 1978b). As some of them are labile when
other feathers, more numerously with the exposed to light, they tend to occur in feath-
flight feathers than elsewhere. They are the ers protected from sunlight. This may also
only feathers that lack muscles.
Feather mass averages 6% (r 5 4–8%) of diseases and skin infections, and transmit
body mass for most birds. diseases.
Despite appearance, feathers do not arise
uniformly over the body. Contour feathers Follicle and muscles
are arranged in discrete areas (tracts) sepa- Follicles are cylindrical sockets in the
rated by bare or sparsely feathered zones skin that produce feathers and hold them
(apteria). This layout is as basic and char- tightly around the calamus. Their kerati-
acteristic a feature of all birds as feathers nized lining is surrounded by germinative
themselves. Tracts cover only about one- epidermis and dense collagen with elastic
half the skin area of land birds, more in fibers (Ostmann et al., 1963; Lucas and
waterbirds. Even in species where the tracts Stettenheim, 1972). The outer dermal tis-
seem to cover the body completely (adult sues are vascularized and well supplied
ratites, penguins, screamers), their bound- with general somatic sensory fibers (Stam-
have 1–2 of these receptors and filoplumes calamus is the last. Hormones and other
have more. factors that affect feather growth exert their
Follicle morphology is nearly the same influence in this sequence.
for all kinds of feathers in many kinds of Feather parts begin to keratinize after
birds. Highly conservative, it varies only in their morphological development is com-
size and in feather musculature. A follicle’s plete, again starting at the tips. The process
histology gives no hint of the kind of feath- is similar to that in reptilian epidermis al-
er that will grow from it, let alone the de- though it happens much later. The rachis
tails of that feather. arises on the dorsal side of the feather tube
and the barbs, lengthening basally, grow
Feather growth and replacement around to join it. The shaft and barbs of an
Embryology. Feather rudiments begin to afterfeather, if any, arise on the ventral side
form in an embryo through a series of in- of the tube. Barbules likewise form in place
Brush, A. H. and J. A. Wyld. 1982. Molecular orga- McLelland (eds.), Form and function in birds,
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Buchholz, R. 1991. Older males have bigger knobs: ering of ptarmigan feet. Condor 79:380–382.
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