The properties of bird feathers as converse piezoelectric transducers and as receptors of microwave radiation. II. Bird feathers as dielectric receptors of microwave radiation.

The characteristics of bird feathers as receptors of microwave fields were investigated in the 10- to 16-GHz region. Experiments were conducted coupling the specimen (feather) to a length of waveguide which served, together with other microwave components, as a primary detector. Microwave power radiation patterns were measured both in the presence and in the absence of the specimen. Results indicated a substantial increase in the microwave power collected in the forward direction and a decrease of the radiation pattern beam width when the feather was present. Fruthermore, some experiemental evidence indicated the possibility of inducing piezoelectric effects in the specimen by audiofrequency pulse-modulated microwave fields. These results are important in view of (i) the fundamental role that feathers play in the life of birds and (ii) the influence of environmental factors on bird behaviour.     

Morphometry of auricular feathers of barn owls (Tyto alba)

In all owl species, the facial plumage forms a parabolic dish, the facial ruff, which is most conspicuous in the the barn owl (Tyto alba). The center of the ruff is formed by auricular feathers. Such feathers are also found on the preaural flaps which cover the ear openings, and in the region of the beak. In this study, we compare the different types of auricular feathers of the barn owl with contour feathers from the neck. Auricular feathers are characterised by an open vane structure and fewer barbs as compared to contour feathers. Auricular feathers also have fewer distal and proximal barbules than contour feathers. The open vane of the auricular feather results from an acute angle between the barb and the basis of the barbules, and from the extension of the pennula parallel to the barbs. These reductions are differently expressed in the three different types of auricular feathers investigated here and correspond with their function (protecting the ruff from dust).     

The evolutionary origin of feathers.

Previous theories relating the origin of feathers to flight or to heat conservation are considered to be inadequate. There is need for a model of feather evolution that gives attention to the function and adaptive advantage of intermediate structures. The present model attempts to reveal and to deal with, the spectrum of complex questions that must be considered. In several genera of modern lizards, scales are elongated in warm climates. It is argued that these scales act as small shields to solar radiation. Experiments are reported that tend to confirm this. Using lizards as a conceptual model, it is argued that feathers likewise arose as adaptations to intense solar radiation. Elongated scales are assumed to have subdivided into finely branched structures that produced a heat-shield, flexible as well as long and broad. Associated muscles had the function of allowing the organism fine control over rates of heat gain and loss: the specialized scales or early feathers could be moved to allow basking in cool weather or protection in hot weather. Subdivision of the scales also allowed a close fit between the elements of the insulative integument. There would have been mechanical and thermal advantages to having branches that interlocked into a pennaceous structure early in evolution, so the first feathers may have been pennaceous. A versatile insulation of movable, branched scales would have been a preadaptation for endothermy. As birds took to the air they faced cooling problems despite their insulative covering because of high convective heat loss. Short glides may have initially been advantageous in cooling an animal under heat stress, but at some point the problem may have shifted from one of heat exclusion to one of heat retention. Endothermy probably evolved in conjunction with flight. If so, it is an unnecessary assumption to postulate that the climate cooled and made endothermy advantageous. The development of feathers is complex and a model is proposed that gives attention to the fundamental problems of deriving a branched structure with a cylindrical base from an elongated scale.     

Colourful parrot feathers resist bacterial degradation

The brilliant red, orange and yellow colours of parrot feathers are the product of psittacofulvins, which are synthetic pigments known only from parrots. Recent evidence suggests that some pigments in bird feathers function not just as colour generators, but also preserve plumage integrity by increasing the resistance of feather keratin to bacterial degradation. We exposed a variety of colourful parrot feathers to feather-degrading Bacillus licheniformis and found that feathers with red psittacofulvins degraded at about the same rate as those with melanin and more slowly than white feathers, which lack pigments. Blue feathers, in which colour is based on the microstructural arrangement of keratin, air and melanin granules, and green feathers, which combine structural blue with yellow psittacofulvins, degraded at a rate similar to that of red and black feathers. These differences in resistance to bacterial degradation of differently coloured feathers suggest that colour patterns within the Psittaciformes may have evolved to resist bacterial degradation, in addition to their role in communication and camouflage.     

Cell organization of barb ridges in regenerating feathers of the quail: implications of the elongation of barb ridges for the evolution and diversification of feathers

This ultrastructural study on the regenerating feathers of quail describes the cellular organization of the barb ridges responsible for the ramification of adult feathers. Bilateral symmetry of the barb ridges determines the organization of feather cells into feather branching. The length of the barb ridges, derived from the number of cells associated to form the barbule plates, determines the length of the barbule branching. Long chains of barb cells form long barbs that branch from the rachis with an increase of feather size. Supportive cells function as spacers between the barbule cells. New cells derive from stem cells localized in the collar region of the feather follicle, as indicated from the re‐organization of collar cells into barb ridges (a morphogenetic process inherited from that of embryonic feathers), production of an embryonic type of keratin (feather keratin), permanence of periderm granules (typical embryonic organelles) in barb vane ridge cells. Variations in the process of barb ridge morphogenesis allow the fusion of ridges into a rachis. The differentiation of hooklets contributes to the origin of planar feathers. Separation between rachis and merging barb ridges is by supportive cells, derived from the marginal plates of the barb ridges. Speculations on the evolution and diversification of feathers are presented.     

Resonating feathers produce courtship song

Male Club-winged Manakins, Machaeropterus deliciosus (Aves: Pipridae), produce a sustained tonal sound with specialized wing feathers. The fundamental frequency of the sound produced in nature is approximately 1500 Hz and is hypothesized to result from excitation of resonance in the feathers'' hypertrophied shafts. We used laser Doppler vibrometry to determine the resonant properties of male Club-winged Manakin''s wing feathers, as well as those of two unspecialized manakin species. The modified wing feathers exhibit a response peak near 1500 Hz, and unusually high Q-values (a measure of resonant tuning) for biological objects (Q up to 27). The unmodified wing feathers of the Club-winged Manakin do not exhibit strong resonant properties when measured in isolation. However, when measured still attached to the modified feathers (nine feathers held adjacent by an intact ligament), they resonate together as a unit near 1500 Hz, and the wing produces a second harmonic of similar or greater amplitude than the fundamental. The feathers of the control species also exhibit resonant peaks around 1500 Hz, but these are significantly weaker, the wing does not resonate as a unit and no harmonics are produced. These results lend critical support to the resonant stridulation hypothesis of sound production in M. deliciosus.     

Abnormal feathers of the micromelic syndrome in White pekin ducks.

In an effort to provide further information concerning the pleiotropic effects of the gene mutation responsible for micromelia in White Pekin ducklings, a histological examination was made oq the abnormal feathers associated with the mutation. Abnormalities found in mutant feathers included decreased overall size, absence of prelumulae and prefiloplumulae feathers, an abnoramlly small rhachis with a disproportionally small medulla, thickening of the feather-sheath, and increased abundance of pulp cells. Embryos having the most abnormally developed feathers and the thickest periderm and feather-sheaths. The nature of many of the abnormalities found in mutant feathers suggests a common source in defective embryonic mesoderm.     

The colour of fossil feathers

Feathers are complex integumentary appendages of birds and some other theropod dinosaurs. They are frequently coloured and function in camouflage and display. Previous investigations have concluded that fossil feathers are preserved as carbonized traces composed of feather-degrading bacteria. Here, an investigation of a colour-banded feather from the Lower Cretaceous Crato Formation of Brazil revealed that the dark bands are preserved as elongate, oblate carbonaceous bodies 1-2mum long, whereas the light bands retain only relief traces on the rock matrix. Energy dispersive X-ray analysis showed that the dark bands preserve a substantial amount of carbon, whereas the light bands show no carbon residue. Comparison of these oblate fossil bodies with the structure of black feathers from a living bird indicates that they are the eumelanin-containing melanosomes. We conclude that most fossil feathers are preserved as melanosomes, and that the distribution of these structures in fossil feathers can preserve the colour pattern in the original feather. The discovery of preserved melanosomes opens up the possibility of interpreting the colour of extinct birds and other dinosaurs.     

The evolutionary origin and diversification of feathers

Progress on the evolutionary origin and diversification of feathers has been hampered by conceptual problems and by the lack of plesiomorphic feather fossils. Recently, both of these limitations have been overcome by the proposal of the developmental theory of the origin of feathers, and the discovery of primitive feather fossils on nonavian theropod dinosaurs. The conceptual problems of previous theories of the origin of feathers are reviewed, and the alternative developmental theory is presented and discussed. The developmental theory proposes that feathers evolved through a series of evolutionary novelties in developmental mechanisms of the follicle and feather germ. The discovery of primitive and derived fossil feathers on a diversity of coelurosaurian theropod dinosaurs documents that feathers evolved and diversified in nonavian theropods before the origin of birds and before the origin of flight. The morphologies of these primitive feathers are congruent with the predictions of the developmental theory. Alternatives to the theropod origin of feathers are critique and rejected. Hypotheses for the initial function of feathers are reviewed. The aerodynamic theory of feather origins is falsified, but many other functions remain developmentally and phylogenetically plausible. Whatever their function, feathers evolved by selection for a follicle that would grow an emergent tubular appendage. Feathers are inherently tubular structures. The homology of feathers and scales is weakly supported. Feathers are composed of a suite of evolutionary novelties that evolved by the duplication, hierarchical organization, interaction, dissociation, and differentiation of morphological modules. The unique capacity for modular subdivision of the tubular feather follicle and germ has fostered the evolution of numerous innovations that characterize feathers. The evolution of feather keratin and the molecular basis of feather development are also discussed.     

Amplification success of multilocus genotypes from feathers found in the field compared with feathers obtained from shot birds

Effective DNA extraction methods from bird feathers have facilitated non‐invasive sampling, leading to the suggestion that feathers are a great source for genetic studies. However, few studies have assessed whether all feathers can be used or provide equal numbers of useful templates. In this study, feathers collected in various ways from Red Grouse Lagopus lagopus were examined to establish the quality of DNA extracted. Individual samples were classified into two categories according to whether they were collected from shot birds or found in the field. DNA was extracted from all samples and genotyped at 19 microsatellite loci. PCR products were analysed on a MegaBACE 1000. A total of 93% of the ‘shot’ category produced a genotype that was considered successful (i.e. 15 of 18 loci) and 23% of the ‘collected’ category produced successful genotypes under the same criteria. There was a significant difference between shot and collected samples in genotyping success and the observed number of missing loci. Recommendations and best practices are discussed along with the utility of bird feathers as a source of DNA for population and conservation biology.     

Development and evolutionary origin of feathers

Avian feathers are a complex evolutionary novelty characterized by structural diversity and hierarchical development. Here, I propose a functionally neutral model of the origin and evolutionary diversification of bird feathers based on the hierarchical details of feather development. I propose that feathers originated with the evolution of the first feather follicle-a cylindrical epidermal invagination around the base of a dermal papilla. A transition series of follicle and feather morphologies is hypothesized to have evolved through a series of stages of increasing complexity in follicle structure and follicular developmental mechanisms. Follicular evolution proceeded with the origin of the undifferentiated collar (stage I), barb ridges (stage II), helical displacement of barb ridges, barbule plates, and the new barb locus (stage III), differentiation of pennulae of distal and proximal barbules (stage IV), and diversification of barbule structure and the new barb locus position (stage V). The model predicts that the first feather was an undifferentiated cylinder (stage I), which was followed by a tuft of unbranched barbs (stage II). Subsequently, with the origin of the rachis and barbules, the bipinnate feather evolved (stage III), followed then by the pennaceous feather with a closed vane (stage IV) and other structural diversity (stages Va-f). The model is used to evaluate the developmental plausibility of proposed functional theories of the origin of feathers. Early feathers (stages I, II) could have functioned in communication, defense, thermal insulation, or water repellency. Feathers could not have had an aerodynamic function until after bipinnate, closed pennaceous feathers (stage IV) had evolved. The morphology of the integumental structures of the coelurisaurian theropod dinosaurs Sinosauropteryx and Beipiaosaurus are congruent with the model's predictions of the form of early feathers (stage I or II). Additional research is required to examine whether these fossil integumental structures developed from follicles and are homologous with avian feathers. J. Exp. Zool. (Mol. Dev. Evol.) 285:291-306, 1999.Copyright 1999 Wiley-Liss, Inc.     

Biological monitoring of exposure to ambient lead and cadmium using avian feathers: A study from Northern India

Biological monitoring of exposure to ambient environmental lead and cadmium was performed using feathers of 26 species of birds native to Western Uttar Pradesh in Northern India. The rationale of this study was to address three questions. First, is there any avian species that can be treated as a suitable bioindicator of lead or cadmium present in the environment? Second, do the birds selectively accumulate lead and cadmium in their feathers and exhibit interspecies variation? Third, is there any threat to endangered species of this region from metal pollution? Average concentration of lead in the feathers of selected birds ranged from 3.40 µg/g in parrot to 301.6 µg/g in golden pheasant, whereas cadmium concentration was higher ranging from 40.20 µg/g in red crow to 450 µg/g in blue macaw. A comparison of results made through ANOVA revealed a significant difference in the concentration of lead (df = 25; F = 3965.54) and cadmium (df = 25; F = 8537.27) in their feathers. We hypothesize that feathers of synanthropic birds may be treated as suitable noninvasive tool to monitor the ambient environmental contamination by lead and cadmium. Their accumulation in endangered birds may lead to population decline causing serious ecological disturbances in the region.     

Adaptive allocation of stress-induced deformities on bird feathers

Physiological stress during ontogeny is known to cause abnormalities in keratin structures of vertebrates, but little is known about if and how organisms have evolved mechanisms to reduce the negative effects of these abnormalities. Stress experienced during avian feather growth is known to lead to the formation of fault bars, and thereby to the weakening of feathers because of shortage and slimming of barbules. Here we propose and test a new hypothesis (the 'fault bar allocation hypothesis') according to which birds should have evolved adaptive strategies to counteract this evolutionary pressure. In particular, we predicted and tested the idea that in flying birds, natural selection should have selected for mechanisms to reduce fault bar load on feathers with high strength requirements during flight. Data on the growth of feathers of nestling white storks (Ciconia ciconia) revealed a consistent allocation of more, and more intense, fault bars in innermost than in outermost wing feathers as predicted by our hypothesis. Moreover, the same pattern emerged from feathers of adult storks. We discuss the generality of our results, and suggest avenues for further investigations in this area.     

Bristles before down: A new perspective on the functional origin of feathers

Over the course of the last two decades, the understanding of the early evolution of feathers in nonavian dinosaurs has been revolutionized. It is now recognized that early feathers had a simple form comparable in general structure to the hairs of mammals. Insight into the prevalence of simple feathers throughout the dinosaur family tree has gradually arisen in tandem with the growing evidence for endothermic dinosaur metabolisms. This has led to the generally accepted opinion that the early feather coats of dinosaurs functioned as thermo insulation. However, thermo insulation is often erroneously stated to be a likely functional explanation for the origin of feathers. The problem with this explanation is that, like mammalian hair, simple feathers could serve as insulation only when present in sufficiently high concentrations. The theory therefore necessitates the origination of feathers en masse. We advocate for a novel origin theory of feathers as bristles. Bristles are facial feathers common among modern birds that function like mammalian tactile whiskers, and are frequently simple and hair‐like in form. Bristles serve their role in low concentrations, and therefore offer a feasible first stage in feather evolution.     

Evolution of the morphological innovations of feathers

Feathers are complex assemblages of multiple morphological innovations. Recent research on the development and evolution of feathers has produced new insights into the origin and diversification of the morphological innovations in feathers. In this article, I review and discuss the contribution of three different factors to the evolution of morphological innovations in feathers: feather tubularity, hierarchical morphological modularity, and the co-option molecular signaling modules. The developing feather germ is a tube of epidermis with a central dermal pulp. The tubular organization of the feather germ and follicle produces multiple axes over which morphological differentiation can be organized. Feather complexity is organized into a hierarchy of morphological modules. These morphological modules evolved through the innovative differentiation along multiple different morphological axes created by the tubular feather germ. Concurrently, many of the morphological innovations of feathers evolved through the evolutionary co-option of plesiomorphic molecular signaling modules. Gene co-option also reveals a role for contingency in the evolution of hierarchical morphological innovations.     

The color of greater flamingo feathers fades when no cosmetics are applied

Greater flamingos use cosmetic coloration by spreading uropygial secretions pigmented with carotenoids over their feathers, which makes the plumage redder. Because flamingos inhabit open environments that receive direct solar radiation during daytime, and carotenoids bleach when exposed to solar radiation, we expected that the plumage color would fade if there is no maintenance for cosmetic purposes. Here, we show that the concentrations of pigments inside feathers and on the surface of feathers were correlated, as well as that there was a correlation between the concentrations of pigments in the uropygial secretions and on the surface of feathers. There was fading in color (becoming less red) in feathers that received direct solar radiation when there was no plumage maintenance, but not so in others maintained in darkness. When we controlled for the initial color of feathers, the feathers of those individuals with higher concentration of pigments on the feather surfaces were those that lost less coloration after experimental exposure of feathers to sunny conditions. These results indicate that exposure to sunlight is correlated with the fading of feather color, which suggests that individuals need to regularly apply makeup to be more colorful. These results also reinforce the view that these birds use cosmetic coloration as a signal amplifier of plumage color. This may be important in species using highly variable habitats, such as wetlands, since the conditions experienced when molting may differ from those when the signal should be functional, usually months after molting.     

The early evolution of feathers: fossil evidence from Cretaceous amber of France

The developmental stages of feathers are of major importance in the evolution of body covering and the origin of avian flight. Until now, there were significant gaps in knowledge of early morphologies in theoretical stages of feathers as well as in palaeontological material. Here we report fossil evidence of an intermediate and critical stage in the incremental evolution of feathers which has been predicted by developmental theories but hitherto undocumented by evidence from both the recent and the fossil records. Seven feathers have been found in an Early Cretaceous (Late Albian, ca 100 Myr) amber of western France, which display a flattened shaft composed by the still distinct and incompletely fused bases of the barbs forming two irregular vanes. Considering their remarkably primitive features, and since recent discoveries have yielded feathers of modern type in some derived theropod dinosaurs, the Albian feathers from France might have been derived either from an early bird or from a non-avian dinosaur.     

The morphology of neoptile feathers: Ancestral state reconstruction and its phylogenetic implications

Avian neoptile feathers are defined as the first feather generation, which covers the chick after hatching, and usually described as simple structures consisting of numerous downy barbs which are radially symmetrically arranged and come together in a short calamus. In contrast, in some birds (e.g., Anas platyrhynchos, Dromaius novaehollandiae) the neoptile feathers have a prominent rhachis, and therefore display clear bilateral symmetry. Because the symmetrical variety found in neoptile feathers is poorly understood, their morphology was studied in a more comprehensive and phylogenetic approach. Neoptile body feathers from over 22 bird species were investigated using light microscopy, SEM, and MicroCT. Characters such as an anterior–posterior axis, a central rhachis, medullary cells, and structure of the calamus wall were defined and mapped onto recent phylogenetic hypotheses for extant birds. It can be shown that bilaterally symmetric neoptile feathers (with a solid calamus wall) were already present in the stem lineage of crown‐group birds (Neornithes). In contrast, simple radially symmetric neoptile feathers (with a fragile calamus wall) are an apomorphic character complex for the clade Neoaves. The simple morphology of this feather type may be the result of a reduced period of development during embryogenesis. To date, embryogenesis of neoptile feathers from only a few bird species was used as a model to reconstruct feather evolution. Because this study shows that the morphology of neoptile feathers is more diverse and even shows a clear phylogenetic signal, it is necessary to expand the spectrum of “model organisms” to species with bilaterally symmetric neoptile feathers and compare differences in the frequency of feather development from a phylogenetic point of view. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

A biotechnological process for treatment and recycling poultry feathers as a feed ingredient

A strain of Kocuria rosea with keratinolytic capacity was cultured aerobically on submerged feathers to obtain a fermented feather meal (FFM). This FFM enriched with cells of K. rosea mainly contains crude protein (71%). The pepsin digestibility of the fermented product (88%) was similar to the value of the commercial feather meal and more than 70% greater that untreated feathers. The bacterial biomass improved the content of amino acids lysine (3.46%), histidine (0.94%) and methionine (0.69%). Additionally, the amino acid availability tested by in vivo assay was greater than commercial feather meal. The microbial cells also supplied carotenoid pigments to FFM (68 ppm). These results suggest that feather meal enriched with K. rosea may be useful in animal feeding as protein and pigment source.     

How are feathers digested by raptors?

In order to determine the cause of the evident degradation of feathers from ingested prey in pellets regurgitated by raptors, in vitro digestions of whole feather barbs by pellet extracts, pepsin or trypsin were carried out. The material was analysed by using biochemical and electron microscopic methods. The results show that the changes in the feathers which occur in the stomach of the Falconidae do not arise from digestion of keratin but from hydrolysis of protein acting as a cement matter in the feather.