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A review of color-producing mechanisms in feathers and their influence on

preventive conservation strategies

Article  in  Journal of the American Institute for Conservation · February 2014

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 A REVIEW OF COLOR-PRODUCING MECHANISMS IN
FEATHERS AND THEIR INFLUENCE ON PREVENTIVE
CONSERVATION STRATEGIES
RENÉE RIEDLER, CHRISTEL PESME, JAMES DRUZIK, MOLLY GLEESON, AND
ELLEN PEARLSTEIN

Weltmuseum Wien, Neue Burg

Getty Conservation Institute

University of Pennsylvania Museum of Archaeology and Anthropology

Information Studies and UCLA/Getty Master’s Program in Archaeological and
Ethnographic Conservation

Lighting policies as part of a preventive strategy for feather collections in museums often do not consider the differ-
ent colorant systems found in the feathers. Ornithological research about plumage coloration, while targeting an
understanding of signaling in bird behavior, provides valuable information about different colorant systems and
their response to light exposure. Because photosensitivity varies for different colorants, it is important for conser-
vators to be able to identify colorants as a first step in designing a preventive strategy for feathers. Visual assessment
methods are described as a means for helping conservators distinguish colorant mechanisms. Color measurement
methods, especially as used to monitor color change, must be able to reproducibly measure feather colors despite
their complex geometries. Measurement methods are referenced that permit reproducibility by accounting for the
variations in measurement angles and feather position. Studies reviewed indicate that pigment-based color and
structural color do not have the same light sensitivity, with pigment color being more sensitive. This light sensitivity
will depend on the nature, concentration, and chemical environment of the pigment. Recent work provides evidence
that the insensitivity of structural colors has been over-simplified and photochemical effects beyond fading are
necessary and important directions for further study.
KEYWORDS: Feather, Biopigment, Structural color, Iridescent, Scattering, Objects speciality, Risk management,
Accelerated aging, Colorimetry

. INTRODUCTION
Developing lighting policies requires knowledge of
the range of sensitivities for an object or a collection.
Feathers are usually categorized as light sensitive,
based largely on past and current observations of
color changes by curators and conservators (Pearlstein
). The current guidelines for feathers expressed
as ISO blue wool sensitivities are highly conjectural,
but move feathers into a category of medium sensitivity.
They are considered generally as ISO BW - (Ashley-
Smith ) or ISO Blue Wool - (Michalski ,
; Solajic et al. ). This gives a maximum
range of  million lux hours to about  million lux
hours if ultraviolet is filtered before a perceptible
color change is evident.
This information is useful for determining lighting pol-
icies for feathered collections, but there is little guidance
about where specific feathers may fall in that continuum.
Based on ornithological research, it is now known that
feathers derive their colors from different sources,
which Hudon brought to the attention of the museum
conservation community (). It is important to note
that the light sensitivity of a feather is due in part to its
specific coloration system. Therefore, for items of feath-
erwork, the most sensitive feather will determine the
light sensitivity of the whole item. In order to determine
the sensitivities of the feathers on an object, it is impor-
tant that the feather types and their respective sensi-
tivities have been identified. Based on this knowledge,
a plan can then be developed and exhibition duration
and intensity of illumination defined. This approach sim-
plifies documentation and monitoring of an item’s exhi-
bition history in the most accessible and efficient
manner, a tradition not well developed for anthropologi-
cal collections. This paper aims to outline the color-
producing mechanisms in feathers, how these different

© American Institute for Conservation

of Historic and Artistic Works 
DOI: ./Y. Journal of the American Institute for Conservation , Vol.  No. , –
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

types may be discriminated, assessed, and measured, and
the light sensitivities associated with these color-producing
mechanisms, in order to assist conservators in further
understanding feathers, feather color, and the light
sensitivity of feathers.
. COLOR-PRODUCING MECHANISMS IN FEATHERS
. TERMINOLOGY
Before proceeding further it is important to be absol-
utely clear about how color in feathers is produced and
to clarify some of the terminology. It must be under-
stood that this is an active area of research and concepts
are continually being refined and extended.
Birds see in a range of electromagnetic energy that is
different than human vision. Even though the term
“color” essentially defines human perception, this
paper shall use “color” to define perceptions visible to
both birds and humans, expanding the range to –
 nm. The rationale for including color visible to
birds is that the loss of color rendering aspects of feath-
ers, even if visible only to birds, is significant when
assessing photochemical change in feathers.
Feathers owe their color to pigments (often referred
to as biopigments) and to pigments combined with
structurally induced backscattered coloration (often
referred to as structural color). Further, structural
colors exist on a continuum between those created by
randomly located light scattering structures which
produce incoherent scattering and those created by
small-scale ordered structures that produce interference
colors, or coherent scattering. Figure , from Prum and
Torres (), diagrammatically shows the meaning of
coherent versus incoherent backscattering. The differ-
ence between incoherent scattering and coherent scat-
tering is not a sharp distinction (Shawkey et al. )
and a number of quasi-ordered systems combining
aspects of both are possible (Andersson ; Noh
et al. ). For simplicity this paper shall describe
color as if the two forms of structural color are
separate.
Some biopigments exhibit absorbance under UV illu-
mination, appearing dark, drab, or in low contrast
when observed by humans, while others exhibit fluor-
escence, making regions appear brighter, more colorful,
or in high contrast. Fluorescence is defined by the
capacity of a molecule to absorb electromagnetic radi-
ation at one wavelength and immediately emit it as
luminescence at a longer wavelength. Birds are
thought to use fluorescence for signaling. When exam-
ining light-induced damage, wavelengths below 

FIG. . Comparison of incoherent and coherent scattering mechanisms of biological structural color production (after Prum
and Torres ). (a) Incoherent scattering is differential scattering of wavelengths by individual light scattering objects. The
phase relationships among light waves scattered from different objects are random. (b) Coherent scattering is differential
interference or reinforcement of wavelengths scattered by multiple light-scattering objects (x, y). Coherent scattering of
specific wavelengths is dependent on the phase relationship among the scattered waves. Scattered wavelengths that are out of
phase will cancel out, but scattered wavelengths that are in phase will be constructively reinforced and coherently scattered.
Phase relationships of wavelengths scattered by two different objects (x, y) are given by the differences in path lengths of light
scattered by the first object (x, -′) and a second object (y, -′) as measured from the planes perpendicular to the incident
and reflected waves (a, b) in the average refractive index of the medium.

Journal of the American Institute for Conservation , Vol.  No. , –
 RENÉE RIEDLER ET AL.
nm are useful to estimate changes in fluorescence as an
indicator of chemical change.
In summary, the perceived color in a feather, the
degree to which a stimulus can be described as similar
to or different from red, green, blue, and yellow,
depends on the specific combination and respective
weight of the two color-producing mechanisms: one
based on selective absorption of the incident light, the
other on its selective scattering.
While colors resulting from primarily pigment
absorption mechanisms have been well characterized
in a small set of species, researchers have more recently
demonstrated that selective scattering by the feather
structure influences color rendition in these pigmented
feathers. Similarly, feathers whose color is predomi-
nantly created by structural scattering also often
include pigmentary components. In terms of this
study, the only feathers whose colors are purely struc-
tural are those which are pure white. Other authors cat-
egorize feather coloration (i.e. pigment versus
structural) according to the prominent mechanism, so
it is important to note the definitions being assigned
for this paper. Table  provides examples of feathers
from bird species represented in cultural artifacts
from different regions of the world organized according
to color-producing systems.
. FEATHER STRUCTURE
Feathers are the most complex avian epidermal
appendages (Lucas and Stettenheim ). They grow
from their base—the feather follicle—and develop
their branched structure through helical growth of the
barb ridges (Prum and Scott ) (fig. ). Therefore,
the most mature and worn/exposed feather part is
found at the tip of the feather vane. After maturation
no blood vessels connect the feather to the bird organ-
ism, therefore unlike skin, no repair mechanism exists
in feathers in response to damage. However, the
whole feather will be replaced completely once or
twice a year during molt.
. PIGMENTED COLORS
Pigments selectively absorb and reflect wavelengths
of incident light reaching the material surface. In the
case of feathers these colorants are named biopigments:
they are either endogenous (metabolized by the organ-
ism) or exogenous (derived through dietary pathways).
The pigmentation is rarely uniform across the surface
and can vary from dark to light, can occur in combi-
nations of multiple biopigments, and can be incorpor-
ated discretely into the shaft, barbs, and barbules.
Different types of biopigments include carotenoids,
melanin, psittacofulvins, and porphyrins.
Journal of the American Institute for Conservation , Vol.  No. , –
.. CAROTENOIDS
Carotenoids are responsible for red, yellow, and
orange coloration in the feathers of many species. In
birds that have been studied to determine their color-
ation, carotenoids have been biochemically character-
ized in the plumage of roughly  species spanning
seven orders (McGraw a). This is a small portion
of the , species of known birds. Carotenoids are
obtained through diet and are deposited in lipoid dro-
plets in the cells of the feather follicle (Brush ).
After keratinization, i.e., as the feather matures
forming a tough fibrous structure, the lipid solvent dis-
appears and the carotenoid pigment is dispersed in the
keratin matrix (Lucas and Stettenheim ). All caro-
tenoids absorb light in the violet-blue region between
 and  nm (Shawkey and Hill ; Andersson
and Prager ). The reflectance spectrum shows a
typical sigmoid shape (Andersson and Prager )
with a small UV peak (McGraw et al. ). The red
colorants tend to be the unsaturated hydrocarbon
form—carotene. Those containing oxygen and with
less hydrophobic properties—the xanthophylls—are
mainly responsible for the yellow coloration. No
mention of in vivo fluorescence was found but it has
been found that carotenoids can be fluorescent in
vitro (Frank et al. ).
Carotenoids can degrade by oxidation or isomeriza-
tion (Britton ; Doucet and Hill ). Carotenoid
pigments in bird feathers are most often found in the
all-trans form (Britton ; McGraw a) and
exposure to light, heat, oxygen, humidity, or acidity
can promote their isomerization and change in color.
Light can also induce auto-oxidation which breaks
down double-bonded carotenoids and results in
pigment bleaching, i.e. loss of color (Test ;
Johnson and Jones ). But in a feather, the keratin
structure serves to inhibit oxidation, so that carotenoids
are somewhat protected by their proteinaceous
environment. Alternatively, it has been suggested that
carotenoids, especially astaxanthin (Miki ), are
effective antioxidants and therefore protect the
keratin (Krinsky ). While in plants it is well
known that carotenoid protects chlorophyll from
oxidation (Krinsky ), in feather their protective
role has not been convincingly proven in vivo (Britton
).
.. MELANIN
Melanin is one of the most common colorants in
feathers. Melanization is not dependent on diet.
Melanin is responsible for brown, black, buff, and
some reddish brown coloration in feathers. Melano-
somes, granules of proteins cluster-linked to melanin
(Riesz ), are deposited in the epithelial cell of the
feather. The proteinaceous matrix determines their
size and shape, which may occur as sticks, rods,
 TABLE . EXAMPLES OF FEATHERS FROM BIRD SPECIES REPRESENTED IN CULTURAL ARTIFACTS FROM DIFFERENT REGIONS OF THE WORLD AND THEIR RESPECTIVE
a
COLOR-PRODUCING SYSTEM

COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES

Color Producing Species Location/Color of Feathers Example of Cultural Artifact
System
Pigment-based Carotenoids Northern Flicker (Colaptes auratus) Red, yellow color on vanes, shaftsb Flicker headband (California)c
color Pileated Woodpecker (Dryocopus pileatus) Red color on head feathersb Head plume (California)c
Toco Toucan (Ramphastos toco) Red color on feathersc Headdress (Amazon)d
Roseate Spoonbill (Platalea ajaja) Red/pink color on feathersb Feather shield (Aztec)e
Melanin Golden Eagle (Aquila chrysaetos) Brown color on feathersf Dance cape (California)c
Bald Eagle (Haliaeetus leucocephalus) Brown color on feathersf Headdress (Northwest Coast)g
Double-crested Cormorant (Phalacrocorax auritus) Black color on feathersf Parka (Alaska)h
Double-Wattled Cassowary (Casuarius casuarius) Black color on feathersi Hair ornament (New Guinea)j
Psittacofulvins Amazon parrot (Amazona spp.) Red, orange, yellow color on feathersj Headdress (Amazon)j
Porphyrins Great Horned Owl (Bubo virginianus) Red-brown color on feathersj Plume stick (California)c
Iridescent Indian Peafowl (Peacock) (Pavo cristatus) Blue-green color on feathersl Hat (Korea)m
structural Mallard Duck (Anas platyrhynchos) Green color on head/neck feathersl Feathered basket (California)n
color Hummingbird (Trochilidae spp.) Various colors on throat feathersl Pendant triptych (Mexico)o
Journal of the American Institute for Conservation , Vol.  No. , –

Male Raggiana Bird of Paradise (Paradisea raggiana) Green color on throat feathersl Pelt (Papua New Guinea)p
Non-iridescent Steller’s Jay (Cyanocitta stelleri) Blue color on feathersq Headdress (California)r
structural color Blue-and-Yellow Macaw (Ara ararauna) Blue color on feathersl Headdress (Amazon)s
Cotinga (Cotinga spp.) Blue color on feathersl Feather shield (Aztec)e
Kingfisher (Halcyon spp.) Blue/green color on feathersl Headdress (China)t
a
This is not an exhaustive list, and the species/specific feathers in this table are limited to those whose colorant systems have been identified.
b
McGraw (a).
c
Jacknis ().
d
American Museum of Natural History, Eastern Tukanoan headband, Catalog number /.
e
Riedler ().
f
McGraw (b).
g
Burke Museum of Natural History, West Coast Raven headdress, Object number ./.
h
Smithsonian Institution, St. Lawrence Island Yup’ik birdskin parka, NMAI Catalog number ..
i
American Museum of Natural History, New Guinea hair ornament, Catalog Number ./.
j
McGraw (c).
k
National Museum of the American Indian, Parakatêjê headdress, Catalog number /.
l
Prum ().
m
Peabody Essex Museum, Chorip (rush hat), Accession Number E.
n
Bibby ().
o
The Metropolitan Museum of Art, Mexican pendant triptych, Catalog number ...
p
American Museum of Natural History, New Guinea Bird of Paradise pelt, Catalog No: ./.
q
Shawkey and Hill ().
r
National Museum of the American Indian, Hupa headdress, Catalog number /.


s
National Museum of the American Indian, Tapirapé Thunder ceremony headdress, Catalog number /.
t
Salzman Chase and McCarthy ().
 RENÉE RIEDLER ET AL.
FIG. . Structure of pennaceous contour feather.
platelets, and air-filled tubes (Durrer ). The organ-
ization of melanosomes within the keratin cortex is
variable, from highly organized to random.
Melanin exhibits a unique broadband absorbance
spectrum in the UV and visible range (Riesz ).
All melanized feathers contain both eumelanin and
phaeomelanin granules. The first type has large, dark,
and regularly shaped granules while in the second
type the granules are smaller and irregularly shaped
(Siefferman and Hill ). It is mainly the ratio of
both types of melanin rather than the concentration
of each that determines the color of the structure
(McGraw b). Phaeomelanins appear pale brown
to buff and absorb more strongly in the longer wave-
lengths of the visible spectrum due to the presence of
fewer functional (carbonyl) groups, while eumelanins
Journal of the American Institute for Conservation , Vol.  No. , –
with more functional groups appear dark brown to
black (McGraw b). It is unclear if eumelanins
are more susceptible to light-induced degradation
than phaeomelanins (Doucet and Hill ) or if they
have the same sensitivity to visible light (Lee )
with phaeomelanins being more sensitive to UV radi-
ation than eumelanins (Hoting et al. ). At high
humidity, the photodegradation of melanin inside
feathers is affected by both visible and UV radiation,
while at very low humidity, visible wavelengths can
have a more photodegrading power than UVA wave-
lengths (Sharma ).
Studies by various authors (Burtt ; Bonser and
Purslow ; Keyser and Hill ; Ward et al.
; Burtt and Ichida ; Delhey et al. ) have
shown that in addition to ornamental functions,
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES
melanins—and especially eumelanins—provide struc-
tural support to feathers. Inclusion of melanin granules
increases the thickness as well as the hardness of
keratin. Both result to increase the resistance of mela-
nized plumage towards mechanical damage due to
wear or to microorganism activity (Delhey et al.
). Some studies argue that melanin also provides
feather structure with protection from UV (Burtt
; Ward et al. ) and from oxidative stress
(Test ; Delhey et al. ). While melanin is gener-
ally considered an extremely stable molecule, it has
been shown that the physical and structural properties
of eumelanin may be strongly affected through the
removal of bound water which is structurally important
in eumelanin, and may increase its susceptibility to
auto-oxidation (Riesz ). In vitro, melanosomes
show sensitivity to protein-degrading agents (Riesz
). UV radiation has been reported as initiating
degradation of biological membranes by inducing
alteration of proteins and/or oxidation of lipids (Koche-
var ). Two UV-induced processes altering proteins
seem to occur: one is oxygen-dependent and produces
cross linking of proteins; the second one is oxygen inde-
pendent and induces protein loss (Kochevar ).
.. PSITTACOFULVINS
The brilliant red, orange, and yellow color of parrots
is produced by psittacofulvins and restricted to the
order Psittaciformes, which includes parrots, cocka-
toos, lories, and lorikeets. It is quite recent that these
pigments, detected by Krukenberg in  (Krukenberg
) were described in-depth biochemically (Veronelli
et al. ; Stradi et al. ; Morelli et al. ;
McGraw and Nogare ). Psittacofulvins do not
depend on diet. They are formed directly in the kerati-
nizing cytoplasm of the cells (Völker ; Driesen
). In the visible range and near UV, the reflectance
curve of psittacofulvin-based red feathers is similar to
that of carotenoid-based red feathers (Masello et al.
), and it is difficult to discriminate between these
colorants spectrophotometrically. Psittacofulvins
fluoresce under UV radiation (McGraw et al. ;
Berg and Bennet ).
.. PORPHYRINS
Porphyrins are present in the feathers of at least 
orders of birds, including owls and bustards (Gill
). Porphyrins produce a range of colors, including
pink, browns, reds, and greens. There are two classes of
porphyrins used as biopigments in feathers: natural
porphyrin and metalloporphyrins. Metalloporphyrins
are porphyrin plus the inclusion of iron or copper: for
example, turacin (McGraw c) is the biopigment
responsible for the red in the bird family of turacos,
and turacoverdin, in the same family, is responsible
for the green. Turacoverdin is the only green pigment
Journal of the American Institute for Conservation , Vol.  No. , –

in birds known to date, as other green feathers result
most often from a structural blue and a yellow biopig-
ment occurring together, or from a combination of pig-
ments, for instance yellow xanthophylls with brown
melanin (Lucas and Stettenheim ; McGraw
a).
A characteristic observed on the downy, light-
protected part of owl feathers is an intense fluorescent
pink, which is induced by absorption in the UV. The
UV-visible absorption spectrum of the porphyrins has
a very typical pattern showing a strong absorption
band at about  nm followed by several weaker
absorptions at higher wavelengths (– nm)
(Goldoni ). Except for turacin, which is stable
(Völker ), these pigments are generally photoche-
mically unstable (Gill ) and are thus primarily
found only in new feathers.
.. MATRIX OF KERATIN
All biopigments are embedded within the keratin
matrix that constitutes the mature feather. This
matrix is composed of one of the most chemically and
mechanically durable proteins—beta keratin. It
absorbs light in the green, blue, and UV range (Anders-
son ; Brink and van der Berg ) and at  nm
its absorption reaches % (Andersson ).
Because it is so absorbent in the UV, it is a powerful
protective layer for the biopigments embedded within
it. Feathers consisting of the keratin structure only,
i.e. white feathers, will appear bright white (Prum
; Pohland ) under visible light.
Keratin yellows when exposed to UV radiation and
bleaches when exposed to visible blue light. When
exposed to a light source with a spectrum including
both visible light as well as UV radiation, both bleach-
ing and yellowing reactions occur within the feather: it
is therefore difficult to anticipate what will be the final
results after exposure (Lennox and Rowlands ).
Wool keratin has been more extensively studied and
the complex pattern of fluorescence has been proven
(Smith ). The fluorescence results from the emis-
sion of specific amino acids that constitute the protein
chain and also from internal cross-linking or cross-
linking between different molecules. The fluorescence
is therefore expected to be light-susceptible (Smith
). The specific amino acids in wool keratin differ
from those in feathers.
.. STRUCTURAL COLOR
The second color-producing mechanism involved in
feather color does not rely on wavelength absorption
but instead on wavelengths that are either coherently
or incoherently backscattered by the structure of the
material interacting with the incident light (Prum
a, ). Colors resulting from this mechanism
 RENÉE RIEDLER ET AL.
are referred to as structural colors. They can be further
subdivided into two categories, the non-iridescent
structural colors produced in the barbs, such as the
blue feathers in parrots, and the iridescent structural
colors, from hummingbird or peacock feathers, which
are produced in predominantly flat, rotated melanized
barbules (fig. ).
Iridescent colors display a wide range of appearances
and, by definition, prominently change hue with angle
of observation and/or illumination. In strictly defined
iridescence phenomenon, the hue changes have to be
perceptible under omni-directional illumination that
exists in nature, and not simply be measureable:
indeed the strong directionality of the artificial light
source used during measurement can induce hue
changes which are otherwise not perceptible under
natural lighting conditions (Prum ).
... IRIDESCENT STRUCTURAL COLORS
To produce iridescent colors, a feather has to have a
highly defined structure showing a specific and ordered
organization of components with different alternating
refractive indices. Both the differences in refractive
FIG. . Picture of iridescent color produced by humming-
bird feather barbules.
Journal of the American Institute for Conservation , Vol.  No. , –
indices and their spatial periodicity as well as the rela-
tive thickness of the barbules play a key role in deter-
mining which wavelengths are coherently
backscattered (Osorio and Ham ).
One of the main characteristics of iridescent feathers
is that the color varies depending on the angle between
illumination and observation as well as between the
feather orientation and the plane of examination (see
Sections .. and ..). The change in hue character-
istic of iridescence occurs because the scattered light is
traveling shorter or longer distances through the
layers with increasing or decreasing angle (fig. ).
For a better understanding of structural iridescent
color, a thin film optical model was introduced
(Doucet et al. ; Shawkey et al. ; Maia et al.
). This model is based on the thickness of the
keratin layer and its two associated interfaces (air/
keratin and keratin/melanin). Two variations of this
model are prevalent to explain iridescent colors in
feathers—known as “thin layer of keratin on thick
layer of melanin,” and “thin layer of keratin on thin
layer of melanin” (fig. ). The “thin layer of keratin
on thin layer of melanin” model (model  in fig. )
describes powerful iridescent colors such as the one dis-
played by peacock feathers. In this case the incident
light passes through the melanin layer while also inter-
acting with the melanosomes. Their spatial distribution
and periodicity play a role in the coherent backscatter-
ing of the incident light (Durrer ). It is the reason
why melanin has to be understood not only as a biopig-
ment but also as a structural component. In the “thin
layer of keratin on thick layer of melanin” model
(model  in fig. ) the melanin layer absorbs the incident
light and does not play a role in the coherent scattering.
This model explains the mechanism responsible for
subtle iridescence (Maia et al. ) such as observed
on black feathers of the male Blue-black Grassquit
(Volatinia jacarina) (Maia et al. ). In both cases,
the absorption of melanin plays a role in the final
color of an iridescent feather.
... NON-IRIDESCENT STRUCTURAL COLORS
In non-iridescent colors, the structural element that
backscatters wavelengths is not ordered enough to
produce iridescence and is therefore often qualified as
a quasi-ordered structure. This element, named the
spongy structure, is located in barbs and is constituted
by air vacuoles regularly distributed in keratin (fig. ).
The rendered color is related to the difference
between the refractive indices of air bubbles and
keratin and their respective sizes and spatial distri-
butions (Dyck ; Osorio and Ham ). The
fact that the spongy structure mainly reflects shorter
wavelengths is the reason why blue or violet hues are
most often perceived (Andersson ), with a
maximum peak of reflectance around or slightly
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

FIG. . Schematic representation of the two different thin-films models. Curved arrows represent beams of light. The models
represent the two combinations of layers and interfaces for an upper surface of barbules showing a strong iridescence (model )
or a weak iridescence (model ).

below  nm (Andersson ). Very often the overall
hue results from the combination of the different mech-
anisms, and pigment absorption plays a role. For
example, in the blue feathers of the Steller’s Jay,
melanin absorbs incoherently scattered light entering
the feather barb (Shawkey and Hill ), thereby
enhancing color purity. Green colors are often a combi-
nation of non-iridescent blue structural color associated
with the presence of carotenoids, which absorb light in
the shorter wavelength (blue) part of the visible spec-
trum (Shawkey and Hill ) while reflecting in
regions corresponding to yellow-orange.
. COLORATION IN THE ULTRAVIOLET RANGE
In birds, color appearance is used as an honest signal
of quality by which females can select their mates.
Unlike humans, birds have vision that extends into
the ultraviolet range of – nm, so light reflection
in the UV range is also important for signaling. It is
therefore not surprising that a detectable and some-
times substantial reflectance can be measured between
 and  nm (Pohland ). This overall UV
reflection by feather surface results from the incoherent
back-scattering of UV by the keratin surface combined
eventually with their coherent scattering and reinforce-
ment by the spongy structure (Andersson ; Prum
and Torres ). UV-reflecting plumage colors result
from the balance between back-scattering, which
reinforces UV reflectance, and the UV absorption by
the biopigments. Although a UV reflectance component
is present in many structural colors that reflect in the
visible range, it is rarely found alone (Andersson
). UV reflectance can show iridescence and results
in measurable reflectance in the ultraviolet range.
In some cases fluorescence is involved when a biopig-
ment reemits in the visible range part of the absorbed
UV range energy. Fluorescing varies from fuchsia to
light green and adds even more variety to feather
colors. UV-induced fluorescent plumage has only
been recorded in birds in the order Psittaciformes
(Hausmann et al. ).
. ASSESSING COLOR
There are two complementary ways to assess colors in
feather. Visual examination, while providing crucial

Journal of the American Institute for Conservation , Vol.  No. , –
 RENÉE RIEDLER ET AL.

FIG. . Anatomy of spongy medullary barb tissue (Shawkey et al. ). (a) Image of an eastern bluebird S. sialis (photo by
Mark Liu), showing feathers with blue color created by coherent scattering of light from spongy tissue in feather barbs.
(b) SEM image of the outer surface of blue feather barbs illustrating the dimensions (x, y, z) examined in this paper. (c)
Two-dimensional TEM image of a transverse section through a single feather barb. The x and y planes are illustrated.
Moving inwards, the layers shown are the outer keratin cortex (C), spongy medullary layer (SL) and vacuole (V). Cell boundaries
(dark lines) can be seen in the upper left section of the cross section. The square inset shows a close-up of the spongy layer.
The black spots surrounding the vacuoles are melanosomes.

information concerning the nature of the color-produ- color science approaches do not discriminate between
cing mechanism, does not permit quantitative assess- the various mechanisms but they do provide a reliable
ment of the color surface. Color measurement and way to quantify color and objective tools to

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 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

communicate about it. Color measurement is also the

first required step to monitor the surface color of a
feather or a feathered item.
Combining the two approaches helps inform
decisions concerning the care of a feathered item.
Indeed, it can be expected that color, depending on its
production mechanisms, will not have the same sensi-
tivity to environmental agents and therefore care is
based in part by an understanding of the different color-
ation systems represented by the feathers on an object.
Furthermore, color monitoring gives information con-
cerning the efficiency of the care strategy.

. VISUAL EXAMINATION

Visual examination can be very useful in furthering

the understanding of surface appearance and docu-
menting it. It is important to emphasize the fact that
even if the color-producing mechanisms are fundamen-
tally different with regard to wavelength absorption, it
is almost impossible to directly discriminate visually
between a purely pigmentary color and a color resulting
from the combination of pigment and non-iridescent
structural color. Comparing the appearance changes
of a given surface observed under different conditions
of illumination and/or observation provides crucial
information that can be further compiled to discrimi-
nate between the different color mechanisms respon-
sible for a given color.

.. IN VISIBLE LIGHT

Examining the surface of the feather lying on an
opaque substrate under ambient visible light provides
qualitative information concerning the reflected light.
If the feather appears blue without iridescence, the
color is a non-iridescent structural color as no biopig-
ment has been reported to date as responsible for blue
hue. For green feathers that are not from turacos, the
color mechanism is either non-iridescent structural or
a combination of pigment and non-iridescent struc-
tural. As shown on the figure , it is possible to discrimi-
nate iridescent color by changing the angle between the
incident light and the direction of observation: the hue FIG. . Peacock feather observed under ambient light. The
of the surface will not be the same depending on the observation angle was slightly changed between the two
specific geometry. photos, causing the hue to shift from green in (a) to blue in (b).

.. UNDER UV SOURCE

Observing (with protective goggles) the appearance
of feathers under a long-wave UV source can permit
mapping the presence of biopigment(s). The location
of UV-induced fluorescence on a Great Horned Owl
feather (fig. b) reveals the presence of at least one
fluorescing biopigment, which is a likely porphyrin.
.. ON LIGHT TABLE
Feathers are a translucent material: while part of the
incident light is reflected and another part is absorbed
by the biopigment(s) within the structure, the rest is
transmitted through the structure. For this reason, it
can be informative to observe a feather on a light

Journal of the American Institute for Conservation , Vol.  No. , –
 RENÉE RIEDLER ET AL.

FIG. . Dorsal side of freshly plucked Great Horned Owl feathers. (a) Examined under ambient light. (b) Examined under a UV
source ( nm).

table. The feather will be more or less opaque depend- light table helps to discriminate structural color from
ing on its type. pigmentary color: structural color will not be percepti-
Comparing the color appearance of the same feather ble on the light table. In ambient light, a Western Scrub
in reflected ambient light and in transmitted light on a Jay wing feather appears blue with grey-brown edges

Journal of the American Institute for Conservation , Vol.  No. , –
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

FIG. . Western Scrub Jay feather. (a) Dorsal side in ambient light. (b) Dorsal side on the light table. (c) Ventral side in ambient
light.

on the dorsal side from a combination of incoherent
scattering and melanin pigmentation (fig. a), while
on the ventral side the melanin is the predominant
colorant (fig. c). When observed on a light table,
the dorsal side of a Western Scrub Jay feather
appears to have lost the blue hue that characterizes
its appearance in ambient light (fig. b). In these con-
ditions the dorsal side appears to have the same
grayish color as the ventral side when observed in
ambient light. The grayish color is characteristic of
melanin absorption.
.. DISCRIMINATING THE DIFFERENT COLOR-PRODUCING
MECHANISMS
Recognition of the color-producing mechanisms is
useful in evaluating and ranking which environmental
agents will put a feather or feathered collection at risk.
Indeed, pigmentary colors should be more sensitive to
light exposure than structural colors, while the latter
may be more sensitive to pressure, for instance.
Table  summarizes how the different color-producing
mechanisms can be discriminated by visual
examination.

TABLE . DISCRIMINATION OF COLOR-PRODUCING MECHANISMS BY VISUAL EXAMINATION

Pigmentary Color Non-iridescent Structural Iridescent Structural

Color Color
Nature of color- Absorption Scattering + absorption
Scattering +
producing mechanism absorption
Characteristics of the • Similar colors when Difference of colors when Color depends on
color-producing observed on light table observed on light table and on angle of illumination/
mechanism and on opaque substrate opaque substrate observation
• Possible fluorescence under
UV

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 RENÉE RIEDLER ET AL.
. COLOR MEASUREMENT
Color can be measured using either a spectropho-
tometer or a colorimeter. In the reflectance mode, the
instrument measures the percent of reflected light com-
pared to the incident light, while in the transmittance
mode, it is the percent of light transmitted that is
measured. The resulting curves, plotting %Reflectance
or %Transmittance wavelength by wavelength,
provide unambiguous descriptions of the physical inter-
action between the incident light and the measured item.
An overview of color measurement parameters of feath-
ers and analytical instrumentation was provided by the
authors in an earlier paper (Riedler et al. ).
As the human visual system is not sensitive to all
wavelengths equally, describing color sensation
requires converting the obtained spectra to colorimetric
parameters. Colorimetry locates color in a three-
dimensional space, for example, in the CIELAB color
space, and provides a way to objectively quantify
color appearance and to calculate its change.
.. MEASURING COLOR IN ORNITHOLOGY
It is only recently that the fields of ornithology and
animal behavior have been able to apply sophisticated
approaches to the reproducible characterization of
feather color, such as color measurement in reflectance
mode. A detailed description of quantifying and analyz-
ing plumage color, on bird specimens is published by
Andersson and Prager (). Color measurement
devices and methods were developed to measure
materials with flat and opaque surfaces. For this
reason, their use had to be adapted to three-
dimensional surfaces, such as that of a feather.
Although the effect of measurement geometry on
observed color has been described early on (Land
), it was not until recently that the main par-
ameters that specify and define the geometry of color
measurement (fig. a)—the angle between illumination
and observation axes, the angle between the measured
surface plane and the illumination/observation plane,
and the orientation of the surface in regard to the axis
around which the sample surface can rotate (fig. b)
—were introduced systematically into feather color
measurement (Osorio and Ham ).
In her doctoral thesis and subsequent publications,
Santos (, Santos and Lumeij , and Santos
et al. ) recommended color quantification using
the mathematical model of Bidirectional Reflectance
Distribution Function. This function indicates the
reflectance values produced at various incoming
(illumination) and outgoing (observation) directions
relative to the orientation of the feather. Santos also
established the importance of using multiple-angle
geometries to fully characterize iridescent and struc-
tural plumage color. She proposed measuring surface
Journal of the American Institute for Conservation , Vol.  No. , –
reflectance with multi-angle spectrometry using a fiber
optic illuminator and a detector with an adaptable-
angle fiber holder (fig. ).
.. COLOR MONITORING
While in the field of ornithology the main interest of
color study is to understand the influence of color func-
tion on bird behavior, in conservation it is often to pre-
serve the appearance of the artifact. Preventive
conservation aims to stabilize the condition of an item
or a collection. In many cases, monitoring the color of
the surface of an object is a good method to assess
condition stability in a given item. To achieve this
goal, one needs to be certain that the color change
measured is not induced by inherent variability of the
feather structure or measurement geometry (fig. )
but reflects the color changes induced by the environ-
mental impact.
The need to precisely localize the measured spot on
the feather surface and to systemically report the differ-
ent orientation of the surface between the illumination/
observation plane and the surface plane can be achieved
using a system of double Mylar templates. The data are
not published in this paper, but the authors were able to
achieve reproducible and repeatable measurements
using this method of color monitoring. However, irides-
cent feather patches could not be measured in a repea-
table manner, as also pointed out by Meadows et al.
().
. COLOR CHANGE
. COLOR CHANGE STUDIES IN ORNITHOLOGY
Beginning in the s, biological studies of feather
coloration have been enriched by the application of
reflectance spectrophotometry (Dyck ) and by
the use of scanning electron microscopy to better under-
stand structural iridescent and non-iridescent plumage
color mechanisms (Greenewalt et al. ). The
emphasis has been on understanding the correlation
between color appearance and mating choices and on
the condition-dependent expression of color (Anders-
son ).
Feather being dead structure, its coloration has been
understood for a long time as a static trait (Delhey
et al. ), despite a few early studies focusing on
color change (Bancroft ; Test ; Lucas and
Stettenheim ). As it has been recently demon-
strated that seasonal changes also influence signaling
(Ornborg et al. ), more studies have been dedi-
cated to the variety of factors explaining color differ-
ences between molts. Three hypotheses have been
formulated to explain seasonal changes (McGraw
and Hill ; Doucet and Hill ): degradation
of the feather structure through abrasion and wear;
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

FIG. . Parameters of color measurement geometry. (a) The red circle is the target eye. The illumination direction is normal to
the measured surface while the angle (θ) it creates with the direction of observation can vary. For a given angle θ the feather
surface can rotate around the illumination direction with an angle ω. For a given θ and a given ω, the measured surface can
be tilted with an angle w. (b) Examples of different orientations in which the feather can be positioned during measurement
by rotating around the illumination direction and changing the angle ω. The red circle shows the location of the measurement
and has the same area independent of the rotation angle ω, while the area on the feather surface will change with tilt angle w.

intentional or accidental soiling; and degradation of
the pigments involved. Abrasion can be induced by
physical friction or keratin-degrading activities of
microorganisms (Hill et al. ; Shawkey et al.
). Keratin degradation can affect structural but
also pigmentary colors, the embedded pigments
losing physical protection (Blanco et al. ).
Soiling can result from soil bathing, inducing dirt
accumulation that can alter structural color
(Ornborg et al. ; Delhey et al. ), or from
application of cosmetic agents such as preen oils or
waxes for maintenance or camouflage purposes that
can alter the absorbance spectra (Blanco et al. ;
Delhey et al. ).
Most studies conclude seasonal color changes result
from the combination of at least two factors, which
sometimes work together and sometimes mitigate
each other. For instance, light exposure, especially UV
radiation, is known to impair feather structure by dena-
turing keratin and other proteins that constitute feath-
ers (Burtt ; Blanco et al. ) while it also
protects feathers against bacterial activity which is pro-
moted by darkness (Delhey et al. ). Soiling can
alter structural color (Ornborg et al. ; Delhey
et al. ) while also reducing risk of microorganism
degradation. Also the contributing factors to seasonal
color changes can either promote the same type of
color changes or induce changes in the opposite direc-
tion: for instance, dirt accumulation tends to darken

Journal of the American Institute for Conservation , Vol.  No. , –
 RENÉE RIEDLER ET AL.

FIG. . Multi-angled fiber holder on peacock feather.

the original color while light exposure tends to bleach it . LIGHT AGING STUDIES
(Surmacki a).
.. COLOR-PRODUCING MECHANISMS AND LIGHT
Pigment degradation can result from abrasion and/or
SENSITIVITY
chemical bond breakage induced by light exposure.
The distinction between pigmentary (absorbing) and
Melanin is more stable to light exposure than carotenoids
structural (non-absorbing and scattering) contributions
(Burtt ; Figuerola and Senar ; Delhey et al.
to feather coloration is crucial because light-induced
) and it also strengthens keratin structure: it is there-
degradation is a direct consequence of energy absorp-
fore expected that melanized plumage would show less
tion. Light-induced degradation results when absorbed
change than carotenoid-based plumage with the same
energy induces irreversible chemical changes. There-
exposure (Test ; Johnson and Jones ; Blanco
fore, with regard to light-induced degradation it is
et al. ; Figuerola and Senar ). A literature
expected that pigmentary colors will be more sensitive
review through  of the light stability of carotenoid
than structural colors. While in all cases, the feather
pigments appears in Pearlstein and Keene ().
keratin matrix plays a role in color rendition, it is
Eliason and Shawkey () demonstrated a rapid
only in the case of pure white feathers that the light sen-
and reversible shift towards longer wavelengths of blue
sitivity is equivalent to that of keratin.
iridescent plumage color accompanying swelling of the
keratin induced by an increase in ambient humidity.
.. LIGHT AGING STUDIES IN CONSERVATION
In the field of conservation the fading of feather-
work has been the subject of few studies. A study
investigating parakeet feathers was carried out by
Horie () with a second study conducted by

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 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

FIG. . Reflectance spectra of a given location on peacock feather showing the color measurement variability induced by color
measurement geometry (as illustrated in fig. ): (a) when the observation angle θ varies, and (b) when the rotation angle ω varies.

Solajic et al. (). Horie’s long-term study included
window fading of budgerigar (Melopsittaus undalatus)
feathers with and without UV radiation. He concluded
that the fading behavior and the sensitivity towards
UV energy varied with the feather color. Solajic in col-
laboration with Pretzel () investigated the effect of
UV-filtered light on Amazonian featherwork, including
Blue and Yellow Macaw, Scarlet Macaw, and Toucan.
They found that feathers were of medium sensitivity to
light between ISO BWS  and BWS . Pearlstein and
Keene () conducted a study comparing window
fading with accelerated light aging of the carotenoid-
colored shafts of Red-shafted Flickers (Colaptes
auratus cafer). The authors found the feather shafts
to be of medium sensitivity, closest to ISO BWS ,
and that exposure to ultraviolet radiation caused
increased fading. An extensive study of color and
color change in natural history collections was recently
carried out by Pohland (). He measured color
differences of various hummingbird, parrot, and
bowerbird species’ feathers held under different
storage conditions: specimens held in public exhibition
exposed to light on a daily basis and specimens from
scientific collections kept in the dark. He also studied
the impact of environmental agents, including light
exposure, on UV reflectance and explored the
possible correlation between UV absorption and fluor-
escence and the impact of light exposure on these
phenomena.
From this literature review two trends emerge. First,
pigment-based color and structural color do not have
the same light sensitivity, the former being more

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 RENÉE RIEDLER ET AL.
sensitive. Second, the light sensitivity of pigmentary
color will depend on the nature, concentration, and
chemical environment of the pigment. However,
recent work provides evidence that the insensitivity of
structural colors has been over-simplified (Surmacki
b). It was demonstrated that exposure to sunlight
can damage keratin structure through photo-oxidation,
implying that all types of coloration can be affected.
. CONCLUSION
Preventive conservation approaches require both the
ability to identify the main risks for a given collection
and the ability to ensure that the collection or item’s
condition is stable in a given controlled environment.
Identifying the main risks for a given collection/item
assumes the identification of the main deleterious
agents and evaluating the sensitivity of the item in
response to each of them. The bibliographical review
concerning color changes of feathers produced in the
field of ornithology provides crucial information con-
cerning the different adverse agents with regard to
feathered collections. It also assists in ranking adverse
agents relative to each specific color-producing mech-
anism. Therefore, for a given feather, a better under-
standing of the different mechanisms involved in its
coloration helps to better identify risk and to implement
preventive conservation policies. For instance, pigmen-
tary colors should be more sensitive to light exposure
than structural colors while pressure can put non-
iridescent structural color at higher risk.
Identifying the feather and if possible its color chem-
istry and structure is crucial, though only a small
portion of bird species have identified coloration
systems. More specifically, with regard to light sensi-
tivity assessment, the basic rule of setting the overall
light sensitivity of the item as equal to the most sensitive
part of a surface can be applied, i.e. assessing the light
sensitivity of the most sensitive pigmentary color of
the surface is crucial to set an appropriate lighting
policy. As mentioned earlier, while it is fairly easy to
recognize an iridescent color by just tilting the feather
or by moving the head while looking at it, it is almost
impossible to discriminate between a pigmentary
color and a non-iridescent structural color. However,
as shown in the paper, comparison of color appear-
ances of a given feather under different conditions of
exam can be very helpful in discriminating between
the two. Combining observations made by examining
the feather in ambient light at various angles of obser-
vation or when the surface is lit with a long-wave UV
source or lying on a light table seems particularly
useful.
The proposed method of combining the information
provided by visual examination conducted by changing
the illumination/observation conditions may have to be
Journal of the American Institute for Conservation , Vol.  No. , –
adapted, consolidated, and standardized by the conser-
vator in a conservation lab but it already appears to be
a low-technology, user-friendly, efficient method to dis-
criminate between the different color-producing mech-
anisms. Discriminating between the different color-
producing mechanisms will also help in selecting the
location to monitor color on the feather surface in an
attempt to stabilize its condition. Indeed, stabilizing
the condition of an item is the goal of any preventive
conservation strategy and it is common to assess the
condition of an item by monitoring its color. Monitor-
ing color on feathers requires developing methods that
control and report all the geometrical parameters that
have been demonstrated to affect the measurement.
The authors were able to develop such a color-
monitoring method, adapting tools and methods used
in the ornithological field, as described earlier in this
paper. While technical details and results of the color
monitoring method are not described in this paper, a
publication describing the method is in preparation.
Based on this work, a preventive conservation strategy
may be developed for a feathered collection.
More generally, distinguishing between the scatter-
ing-based mechanism and the absorption-based mech-
anism may be useful in order to assess the risk of
appearance change of a material when exposed to a
specific adverse agent. While feather provides a particu-
larly clear illustration of the respective contributions of
physical and chemical color mechanisms, both mechan-
isms operate to varying degrees in the rendering of the
appearance of any surface. The respective contribution
of the scattering-based and the absorption-based mech-
anisms create the appearance of color, gloss and trans-
parency of a given material and its specific sensitivity
towards a given environmental agent.
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AUTHOR BIOGRAPHIES
RENÉE RIEDLER received an MA in Philology and Art History from the University of Vienna and an MA from the Academy of Fine
Arts in Conservation and Restoration. As postgraduate intern and consultant she was researching full time for the collaborative
feather research project between UCLA and the Getty Conservation Institute. She is currently living and working in Vienna,
Austria. Address: Czerningasse //, A- Vienna, Austria. Email: renee.riedler@hotmail.com

CHRISTEL PESME received a BA in Biochemistry from the University of Paris -Pierre et Marie Curie, an MA in Art History and an
MA in paper conservation, both from the University of Paris -Pantheon Sorbonne (France). She has been working with Jim
Druzik as an assistant scientist at the Getty Conservation Institute, Los Angeles, carrying out light sensitivity for the Getty
Research Institute, and the J. Paul Getty Museum. She is now working in Basel (Switzerland) as a private conservator and
part time on writing her PhD in Art History. Address: Utengasse ,  Basel, Switzerland. Email: Christel.pesme@gmail.com

JAMES DRUZIK is a Senior Scientist at the Getty Conservation Institute, where he has worked since . His research interests have
been highly varied over the years and have included image processing (Caltech/JPL), the origin and fate of oxidant air pollutants

Journal of the American Institute for Conservation , Vol.  No. , –
 RENÉE RIEDLER ET AL.

and particulates in museum environments and their control technologies in conjunction with Glen Cass (Caltech). His group now
routinely carries out assessment of light sensitivity for the Getty Research Institute, and the J. Paul Getty Museum and carries out
research on solid-state lighting. He has previously worked at the Los Angeles County Museum of Art and the Norton Simon
Museum in Pasadena, California. Druzik holds a BS in Chemistry. Address: Getty Conservation Institute,  Getty Center
Drive, Suite , Los Angeles, CA . Email: JDruzik@getty.edu

MOLLY GLEESON received a BA in Art Conservation from the University of Delaware and an MA from the UCLA/Getty Master’s
Program in the Conservation of Archaeological and Ethnographic Materials. She is currently working in San Diego, California,
devoting part of her time to assist on this collaborative feather research project between UCLA and the Getty Conservation Insti-
tute. Address:  Campus Ave., San Diego, CA . Email: mcgleeson@yahoo.com

ELLEN PEARLSTEIN is an associate professor at the University of California, Los Angeles. Her research interests include American
Indian tribal museums and how museum staff defines cultural preservation; effects of environmental agents on ethnographic and
natural history materials; introducing context into cultural materials’ conservation education; and curriculum development. She
is co-director of the UCLA and Getty Conservation Institute feather research project. Address: Information Studies and
UCLA/Getty Master’s Program in Archaeological and Ethnographic Conservation, A Fowler, Los Angeles, CA .
Email: epearl@ucla.edu

Résumé–Souvent dans le cadre d’une stratégie préventive pour les collections de plumes dans les musées, les normes
d’éclairage ne considèrent pas les différents systèmes de colorants présents dans les plumes. La recherche ornitho-
logique sur la coloration du plumage, tout en ciblant la compréhension de la signalisation dans le comportement des
oiseaux, fournit des informations importantes sur les différents systèmes de colorants et leur réaction à l’exposition
à la lumière. Parce que la photosensibilité varie selon les colorants, il est important pour les restaurateurs de pouvoir
identifier les colorants dans une première étape lors de la conception d’une stratégie préventive pour les plumes. Des
méthodes d’évaluation visuelle sont décrites comme moyen pour aider les restaurateurs à distinguer les mécanismes
des colorants. Les méthodes pour mesurer les couleurs, particulièrement pour traquer les changements de couleur,
doivent être capables de mesurer de manière reproductible la couleur des plumes malgré leurs géométries complexes.
Les auteurs font référence à différentes méthodes qui permettent une certaine reproductibilité en tenant compte des
variations dans les angles de mesure et dans la position des plumes. Les études passées en revue indiquent que les
couleurs à base de pigments et les couleurs structurelles n’ont pas la même sensibilité à la lumière, les couleurs à base
de pigment étant plus sensibles. Cette sensibilité à la lumière dépend de la nature, de la concentration et de l’envir-
onnement chimique du pigment. Des travaux récents fournissent des preuves que l’insensibilité des couleurs struc-
turelles a été trop simplifiée et indiquent que les effets photochimiques au-delà de la perte des couleurs doivent
nécessairement devenir le sujet d’études plus approfondies.

Resumen—Las políticas de iluminación que son parte de la estrategia preventiva en las colecciones de plumas en los
museos, con frecuencia no tienen en cuenta los diferentes sistemas colorantes que se encuentran en las plumas. Las
investigaciones ornitológicas sobre la coloración de plumas, al mismo tiempo que se enfocan en entender las señales
del comportamiento de las aves, también nos ofrecen información valiosa a cerca de los diferentes sistemas color-
antes y su respuesta a la exposición a la luz. Ya que la sensibilidad a la luz varía en los diferentes colorantes, es
importante para los conservadores poder identificar los colorantes como un primer paso al diseñar una estrategia
preventiva para las plumas. Se describen métodos para hacer un diagnostico visual que ayude a los conservadores a
diferenciar los mecanismos de coloración. Los métodos de medición de color, especialmente los que se utilizan para
monitorear los cambios de color, deben ser capaces de medir los colores de las plumas en forma reproducible
aunque estas tienen geometrías complejas. Los métodos de medición son referenciados de manera que permiten
ser reproducidos al tener en cuenta las variaciones en ángulos de medida y posición de las plumas. Por medio de
estudios se descubrió que los colores de los pigmentos y los colores estructurales no tienen la misma sensibilidad
a la luz, siendo el color de los pigmentos mas sensible. Esta sensibilidad a la luz depende de la naturaleza, la con-
centración y el entorno químico del pigmento. Los estudios mas recientes ofrecen evidencia de que la insensibilidad
de los colores estructurales ha sido excesivamente simplificada, y que es necesario e importante realizar estudios mas
avanzados sobre los efectos fotoquímicos mas allá de la decoloración por ser esta una dirección necesaria e
importante.

Resumo—Políticas de iluminação como parte de uma estratégia preventiva para coleções de plumagens em museus
normalmente não consideram os diferentes sistemas de corantes encontrados nas penas. Pesquisa ornitológica sobre
coloração de plumagem, ao objetivar uma compreensão da sinalização no comportamento de pássaros, fornece
informação valiosa sobre os diferentes sistemas de corantes e sua resposta à exposição à luz. Devido à variação

Journal of the American Institute for Conservation , Vol.  No. , –
 COLOR-PRODUCING MECHANISMS IN FEATHERS AND THEIR INFLUENCE ON PREVENTIVE CONSERVATION STRATEGIES 

de fotossensibilidade para diferentes corantes, é importante que os conservadores sejam capazes de identificar cor-
antes como um primeiro passo na concepção da estratégia preventiva para penas. Métodos de avaliação visual são
descritos como um meio para ajudar conservadores a distinguirem os mecanismos de corantes. Métodos de medição
de cor, especialmente os usados para monitorar as mudanças de cor, devem ser capazes de reproduzir medidas das
cores das penas apesar das suas complexidades geométricas. Métodos de medição são referenciados de maneira que
permitam a reprodutibilidade pela contagem das variações nas medidas de ângulos e posição das penas. Os estudos
revisados indicam que as cores à base de pigmentos e cores estruturais não tem a mesma sensibilidade à luz, com a
cor do pigmento sendo mais sensível. Essa sensibilidade à luz dependerá da natureza, concentração e ambiente
químico do pigmento. Trabalhos recentes evidenciam que a insensibilidade das cores estruturais tem sido excessi-
vamente simplificada e os efeitos fotomecânicos além do esmaecimento são caminhos importantes para estudo
posterior.

Journal of the American Institute for Conservation , Vol.  No. , –

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