To pluck or not to pluck: the hidden ethical and scientific costs of relying on feathers as a primary source of DNA

This article responds to the recent prominence of ornithological literature advocating the plucking or clipping of feathers to obtain DNA in avian studies. We argue that the practise of feather plucking or clipping should be strongly discouraged on both scientific and ethical grounds in the avian literature. Currently, despite claims to the contrary, it is not clear that feather sampling as a source of DNA has lower ethical impacts on birds than blood sampling. In addition, feather samples provide a smaller and less reliable biological resource, significantly jeopardising the short and long‐term outcomes that can be gained by the sampling. In contrast, blood collection has been experimentally demonstrated to be relatively safe, subject to operators being skilled and following published guidelines, providing large yields of high quality DNA that facilitates archival storage of samples in a manner that the destructive sampling of feathers cannot.     

A comparison of plucked feathers versus blood samples as DNA sources for molecular sexing

ABSTRACT.   Feathers are increasingly collected as a nondestructive source of DNA for avian genetic research. Although feather samples are not optimal in some important ways than more robust blood or tissue samples, feather sampling requires less training for field workers, results in shorter handling times for the organism, generates no hazardous wastes, and requires simpler storage procedures. Along with these largely positive attributes comes a set of challenges, particularly the relatively low copy number of DNA present in feather samples. We compared the utility and reliability of feathers to the more traditional blood samples as sources of DNA for polymerase chain reaction (PCR)-based molecular sexing of Black-capped Chickadees ( Poecile atricapilla ). DNA from 102 individuals was extracted separately from both single rectrices and from blood samples, and the sex of each bird was then determined using standard PCR-based methods. We found complete agreement between sex determinations based on feather versus blood DNA extractions. Slight variations in lab protocols were necessary to obtain consistent results from these two DNA sources; and we briefly discuss other sources of error that could occur in feather-based molecular sexing studies. This controlled comparison of feather versus blood samples demonstrates that plucked rectrices provide a highly reliable source of DNA for molecular sexing of wild birds.     

Trace element contamination in feather and tissue samples from Anna’s hummingbirds

Trace element contamination (17 elements; Be, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, As, Se, Mo, Cd, Ba, Hg, Tl, and Pb) of live (feather samples only) and deceased (feather and tissue samples) Anna's hummingbirds (Calypte anna) was evaluated. Samples were analyzed using inductively coupled plasma-mass spectrometry (ICP-MS; 17 elements) and atomic absorption spectrophotometry (Hg only). Mean plus one standard deviation (SD) was considered the benchmark, and concentrations above the mean + 1 SD were considered elevated above normal. Contour feathers were sampled from live birds of varying age, sex, and California locations. In order to reduce thermal impacts, minimal feathers were taken from live birds, therefore a novel method was developed for preparation of low mass feather samples for ICP-MS analysis. The study found that the novel feather preparation method enabled small mass feather samples to be analyzed for trace elements using ICP-MS. For feather samples from live birds, all trace elements, with the exception of beryllium, had concentrations above the mean + 1 SD. Important risk factors for elevated trace element concentrations in feathers of live birds were age for iron, zinc, and arsenic, and location for iron, manganese, zinc, and selenium. For samples from deceased birds, ICP-MS results from body and tail feathers were correlated for Fe, Zn, and Pb, and feather concentrations were correlated with renal (Fe, Zn, Pb) or hepatic (Hg) tissue concentrations. Results for AA spectrophotometry analyzed samples from deceased birds further supported the ICP-MS findings where a strong correlation between mercury concentrations in feather and tissue (pectoral muscle) samples was found. These study results support that sampling feathers from live free-ranging hummingbirds might be a useful, non-lethal sampling method for evaluating trace element exposure and provides a sampling alternative since their small body size limits traditional sampling of blood and tissues. The results from this study provide a benchmark for the distribution of trace element concentrations in feather and tissue samples from hummingbirds and suggests a reference mark for exceeding normal. Lastly, pollinating avian species are minimally represented in the literature as bioindicators for environmental trace element contamination. Given that trace elements can move through food chains by a variety of routes, our study indicates that hummingbirds are possible bioindicators of environmental trace element contamination.     

Measuring corticosterone in feathers using an acetonitrile/hexane extraction and enzyme immunoassay: feather corticosterone levels of food‐supplemented Atlantic Puffin chicks

Glucocorticoid levels measured in the blood of animals reflect hypothalamic‐pituitary‐adrenal (HPA) activity in response to predictable and unpredictable changes. In birds, circulating corticosterone is incorporated into growing feathers and provides an integrated measure of HPA activity over the period of feather growth. Measuring corticosterone in feathers can provide insight into the physiological state of birds during times when they are unavailable for blood sampling (e.g., during migratory or non‐breeding periods of the annual cycle). Building upon studies that used radioimmunoassay or liquid chromatography, we used a commercially available enzyme immunoassay kit to measure corticosterone in feathers of nestling Atlantic Puffins (Fratercula arctica) on Gull Island, Newfoundland, Canada, in 2012, and demonstrate the benefits of sample preparation via acetonitrile/hexane purification. We used this method to measure corticosterone in feathers of Atlantic Puffin chicks that experienced differences in mass gain in a supplementary feeding study. We found a positive relationship between feather corticosterone and mass gain, and a negative relationship between feather corticosterone and pre‐treatment body condition. Because feathers were growing prior to and during the supplementary feeding period, our results also suggest that extracting seabird feather samples with acetonitrile/hexane (in addition to methanol) prior to measuring corticosterone with enzyme immunoassay is beneficial, and, as reported in previous studies, blood and feather corticosterone values reflect different measures.     

Co‐infections by malaria parasites decrease feather growth but not feather quality in house martin

During moult, stressors such as malaria and related haemosporidian parasites (e.g. Plasmodium and Haemoproteus) could affect the growth rate and quality of feathers, which in turn may compromise future reproduction and survival. Recent advances in molecular methods to study parasites have revealed that co‐infections with multiple parasites are frequent in bird–malaria parasite systems. However, there is no study of the consequences of co‐infections on the moult of birds. In house martins Delichon urbica captured and studied at a breeding site in Europe during 11 yr, we measured the quality and the growth rate of tail feathers moulted in the African winter quarters in parallel with the infection status of blood parasites that are also transmitted on the wintering ground. Here we tested if the infection with two haemosporidian parasite lineages has more negative effects than a single lineage infection. We found that birds with haemosporidian infection had lower body condition. We also found that birds co‐infected with two haemosporidian lineages had the lowest inferred growth rate of their tail feathers as compared with uninfected and single infected individuals, but co‐infections had no effect on feather quality. In addition, feather quality was negatively correlated with feather growth rate, suggesting that these two traits are traded‐off against each other. We encourage the study of haemosporidian parasite infection as potential mechanism driving this trade‐off in wild populations of birds.     

Feather function and the evolution of birds

The ability of feathers to perform many functions either simultaneously or at different times throughout the year or life of a bird is integral to the evolutionary history of birds. Many studies focus on single functions of feathers, but any given feather performs many functions over its lifetime. These functions necessarily interact with each other throughout the evolution and development of birds, so our knowledge of avian evolution is incomplete without understanding the multifunctionality of feathers, and how different functions may act synergistically or antagonistically during natural selection. Here, we review how feather functions interact with avian evolution, with a focus on recent technological and discovery-based advances. By synthesising research into feather functions over hierarchical scales (pattern, arrangement, macrostructure, microstructure, nanostructure, molecules), we aim to provide a broad context for how the adaptability and multifunctionality of feathers have allowed birds to diversify into an astounding array of environments and life-history strategies. We suggest that future research into avian evolution involving feather function should consider multiple aspects of a feather, including multiple functions, seasonal wear and renewal, and ecological or mechanical interactions. With this more holistic view, processes such as the evolution of avian coloration and flight can be understood in a broader and more nuanced context.     

Quantifying structural variation in contour feathers to address functional variation and life history trade‐offs

Body feathers are important to many interactions birds have with their physical and social environments, such as streamlining the body for flight, thermoregulation, and social signaling. Birds differ dramatically in the texture of their body plumage depending on species and age class, likely reflecting different functional demands and age‐related trade‐offs in feather production. Despite the important insights potentially offered by studying variation in the structure of body feathers, there is no clear system for quantifying this variation. We present methods for quantifying age and species differences in the structure of body feathers. Most variation in our measures is due to species and age‐class differences, with little variance attributable to individual birds or to differences among feathers sampled from the same bird. We use our measures to test the hypothesis that the loosely‐textured plumage characteristic of many juvenile passerines reflects a trade‐off between investment in feather quality and rapid body growth that promotes early fledging. The structure of juvenile feathers was negatively correlated with duration of the nestling period among ten species of New World warblers (Parulidae), suggesting a trade‐off between investment in feathers and investment in rapid somatic development promoting fledging. Systematic studies of variation in the structure of body feathers will likely offer numerous other insights into avian biology.     

Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds

Chan, N.R., Dyke, G.J. & Benton, M.J. 2013: Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds. Lethaia, Vol. 46, pp. 146–152. Although many Mesozoic fossil birds have been found with primary feathers preserved, these structures have rarely been included in morphometric analyses. This is surprising because the flight feathers of modern birds can contribute approximately 50% of the total wing length, and so it would be assumed that their inclusion or exclusion would modify functional interpretations. Here we show, contrary to earlier work, that this may not be the case. Using forelimb measurements and primary feather lengths from Mesozoic birds, we constructed morphospaces for different clades, which we then compared with morphospaces constructed for extant taxa classified according to flight mode. Consistent with older work, our results indicate that among extant birds some functional flight groups can be distinguished on the basis of their body sizes and that variation in the relative proportions of the wing elements is conservative. Mesozoic birds, on the other hand, show variable proportions of wing bones, with primary feather length contribution to the wing reduced in the earlier diverging groups. We show that the diverse Mesozoic avian clade Enantiornithes overlaps substantially with extant taxa in both size and limb element proportions, confirming previous morphometric results based on skeletal elements alone. However, these measurements cannot be used to distinguish flight modes in extant birds, and so cannot be used to infer flight mode in fossil forms. Our analyses suggest that more data from fossil birds, combined with accurate functional determination of the flight styles of living forms is required if we are to be able to predict the flight modes of extinct birds. □Birds, flight, morphospace, Mesozoic, wing.     

Linking the molecular evolution of avian beta (β) keratins to the evolution of feathers

Feathers of today's birds are constructed of beta (β)-keratins, structural proteins of the epidermis that are found solely in reptiles and birds. Discoveries of "feathered dinosaurs" continue to stimulate interest in the evolutionary origin of feathers, but few studies have attempted to link the molecular evolution of their major structural proteins (β-keratins) to the appearance of feathers in the fossil record. Using molecular dating methods, we show that before the appearance of Anchiornis (～155 Million years ago (Ma)) the basal β-keratins of birds began diverging from their archosaurian ancestor ～216?Ma. However, the subfamily of feather β-keratins, as found in living birds, did not begin diverging until ～143?Ma. Thus, the pennaceous feathers on Anchiornis, while being constructed of avian β-keratins, most likely did not contain the feather β-keratins found in the feathers of modern birds. Our results demonstrate that the evolutionary origin of feathers does not coincide with the molecular evolution of the feather β-keratins found in modern birds. More likely, during the Late Jurassic, the epidermal structures that appeared on organisms in the lineage leading to birds, including early forms of feathers, were constructed of avian β-keratins other than those found in the feathers of modern birds. Recent biophysical studies of the β-keratins in feathers support the view that the appearance of the subfamily of feather β-keratins altered the biophysical nature of the feather establishing its role in powered flight.     

Nutritional stress affects corticosterone deposition in feathers of Caspian tern chicks

Stressful environmental conditions affect the adrenocortical function of developing animals, which can have consequences for their fitness. Discovery of the avian stress hormone corticosterone (CORT) in feathers has the potential to broaden the application of endocrine research in ecological and evolutionary studies of wild birds by providing a long‐term measure of CORT secretion. Mechanisms of CORT deposition in feathers are not well known and few studies have related feather CORT to circulating plasma CORT during feather growth. Our objective was to experimentally test the validity of using feather CORT as a measure of CORT secretion in developing birds experiencing nutritional stress. Caspian tern Hydroprogne caspia chicks were fed ad libitum or restricted (35% less than ad libitum) diets for four weeks. We measured CORT in feathers from these chicks to examine the relationship between feather CORT concentrations and nutritional limitation, circulating plasma CORT, and feather development. We found that feather CORT was higher in controls fed ad libitum than in restricted individuals, despite higher levels of plasma CORT in restricted chicks compared to controls. Feather mass and growth rates were strongly and positively related to feather CORT concentrations in both treatments. This is the first experimental study to show that feather CORT concentrations can be lower in response to nutritional stress, even when plasma CORT concentrations are elevated. Our results indicate that CORT deposition in feathers may be confounded when feather mass and growth rates are compromised by nutritional stress. We conclude that feather CORT can be used for assessing nutritional stress in growing birds, but the direction of response depends on how strongly stress affects feather development.     

Fine‐tuned distribution of feather mites (Astigmata) on the wing of birds: the case of blackcaps Sylvia atricapilla

The distribution of feather mites (Astigmata) along the wing of passerine birds could change dramatically within minutes because of the rapid movement of mites between feathers. However, no rigorous study has answered how fine‐tuned is the pattern of distribution of feather mites at a given time. Here we present a multiscale study of the distribution of feather mites on the wing of non‐moulting blackcaps Sylvia atricapilla in a short time period and at a single locality. We found that the number and distribution of mites differed among birds, but it was extremely similar between the wings of each bird. Moreover, mites consistently avoided the first secondary feather, despite that it is placed at the centre of the feathers most used by them. Thus, our results suggest that feather mites do precise, feather‐level decisions on where to live, contradicting the current view that mites perform “mass”, or “blind” movements across wing feathers. Moreover, our findings indicate that “rare” distributions are not spurious data or sampling errors, but each distribution of mites on the wing of each bird is the outcome of the particular conditions operating on each ambient‐bird‐feather mite system at a given time. This study indicates that we need to focus on the distribution of feather mites at the level of the individual bird and at the feather level to improve our understanding of the spatial ecology of mites on the wings of birds.     

Exogenous and endogenous corticosterone in feathers

In birds, the steroid hormone corticosterone (CORT) increases in response to real or perceived threats to homeostasis. A long‐term record of CORT exposure is recorded in feathers when the hormone is incorporated into the keratinized tissue, and then preserved when the mature feather is cut off from the blood supply. The opportunity to retrospectively assess the exposure of an individual to stressors by measuring the amount of CORT in a feather has generated excitement amongst avian ecologists. However, this technique is relatively new and requires additional validations. In this study, we performed experiments in wild caught European starlings Sturnus vulgaris to test whether: 1) CORT deposition in the feather depends on time of day and 2) whether an ecologically relevant stressor (unpredictable food access) causes a change in feather CORT. We found that exogenous CORT was incorporated into feathers during the day and the night. However, there was no difference in feather CORT between birds with unpredictable access to food and those with continuous access, indicating that feather CORT might not always detect ecologically relevant stressors.     

The scaling of the size and stiffness of primary flight feathers

Birds encompass a large range of body sizes, yet the importance of body size on feather morphology and mechanical properties has not been characterized. In this study, I examined the scaling relationships of primary flight feathers within a phylogenetically diverse sample of avian species varying in body size by nearly three orders of magnitude. I measured the scaling relationships between body mass and feather linear dimensions as well as feather flexural stiffness. The resnlts of an independent contrasts analysis to test the effects of phylogenetic history on the characters measured had no effect on the scaling relationships observed. There was slight, but not significant, positive allometry in the scaling of shaft diameter with respect to feather length across a range of body masses. The scaling of feather length and diameter against body mass was not significantly different from isometry. Flexural stiffness, however, exhibited strong negative allometry. Therefore, larger birds have relatively more flexible feathers than smaller birds. The more flexible primary feathers of large birds may reduce stresses on the wing skeleton during take-off and landing and also make these feathers less susceptible to mechanical failure. Conversely, the greater flexibility of these feathers may also reduce their capacity to generate aerodynamic lift.     

Variations in the morphology,distribution, and arrangement of feathers in domesticated birds

Domesticated birds exhibit a greater diversity in the morphology of their integument and its appendages than their wild ancestors. Many of these variations affect the appearance of a bird significantly and have been bred selectively by poultry and pigeon fanciers and aviculturists for the sake of visual appeal. Variations in feather distribution (e.g., feathering of legs and feet, featherless areas in normally feather-bearing skin) are widespread in chickens and pigeons. Variations in the number of feathers (e.g., increased number of tail feathers, lack of tail feathers) occur in certain pigeon and poultry breeds. Variations in feather length can affect certain body regions or the entire plumage. Variations in feather structure (e.g., silkiness, frilled feathering) can be found in exhibition poultry as well as in pet birds. Variations in feather arrangement (e.g., feather crests and vortices) occur in many domesticated bird species as a results of mutation and intense selective breeding. The causes of variations in the structure, distribution, length and arrangement of feathers is often unknown and opens a wide field for scientific research under various points of view (e.g., morphogenesis, pathogenesis, ethology, etc.). To that extent, variations in the morphology, distribution and arrangement of feathers in domesticated birds require also a concern for animal welfare because certain alleles responsible for integumentary variations in domesticated birds have pleiotropic effects, which often affect normal behaviour and viability.     

Malaria infection negatively affects feather growth rate in the house sparrow Passer domesticus

Birds often face various stressors during feather renewing, for example, enduring infection with blood parasites. Because nutritional resources are typically limited, especially for wild animals, when an individual allocates energy to one physiological system, there is subsequently less for other processes, thereby requiring a trade‐off. Surprisingly, potential trade‐offs between malaria infection and feather growth rate have not been experimentally considered yet. Here, we conducted three studies to investigate whether a trade‐off occurs among feather growth rate, malaria infection and host health conditions. First, we explored whether naturally infected and uninfected house sparrows differed in feather growth rate in the wild. Second, we asked whether experimental inoculation of malaria parasites and/or forcing the renewal of a tail feather. Lastly, we evaluated whether individual condition was affected by experimentally‐induced feather regrowth and/or malaria experimental infection. Our findings showed that feather growth rate was negatively affected by natural malaria infection status in free‐living birds and by experimental infection in captive birds. Furthermore, birds that did not increase body mass or hematocrit during the experimental study had slower feather growth. Together our results suggest that infection with blood parasites has more negative health effects than the growth of tail feathers and that these two processes (response to blood parasite infection and renewal of feathers) are traded‐off against each other. As such, our results highlight the role of malaria parasites as a potential mechanism driving other trade‐offs in wild passerines.     

Feather mites (Acari: Astigmata) and body condition of their avian hosts: a large correlative study

Feather mites are arthropods that live on or in the feathers of birds, and are among the commonest avian ectosymbionts. However, the nature of the ecological interaction between feather mites and birds remains unclear, some studies reporting negative effects of feather mites on their hosts and others reporting positive or no effects. Here we use a large dataset comprising 20 189 measurements taken from 83 species of birds collected during 22 yr in 151 localities from seven countries in Europe and North Africa to explore the correlation between feather mite abundance and body condition of their hosts. We predicted that, if wing‐dwelling feather mites are parasites, a negative correlation with host body condition should be found, while a mutualistic interaction should yield positive correlation. Although negative relationships between feather mite abundance and host body condition were found in a few species of birds, the sign of the correlation was positive in most bird species (69%). The overall effect size was only slightly positive (r =0.066). The effect of feather mite abundance explained <10% of variance in body condition in most species (87%). Results suggest that feather mites are not parasites of birds, but rather that they hold a commensalistic relationship where feather mites may benefit from feeding on uropygial gland secretions of their hosts and birds do not seem to obtain a great benefit from the presence of feather mites.     

Feather isotope analysis discriminates age-classes of Western, Least, and Semipalmated sandpipers when plumage methods are unreliable

ABSTRACT Avian age‐class discrimination is typically based on the completeness of the first prebasic molt. In several calidrid sandpiper species, juvenal flight feathers grown on Arctic breeding grounds are retained through the first three migrations. Thereafter, flight feathers are grown annually at temperate migratory stopover sites during the fall or on the subtropical wintering grounds. Standard methods for distinguishing age classes of sandpipers rely on a combination of traits, including body plumage, coloration of protected inner median covert edges, and extent of flight feather wear. We tested the ability of stable hydrogen isotope ratios in flight feathers (δD_f) to distinguish young birds in their first winter through second fall from older adults in three calidrid sandpiper species, Western (Calidris mauri), Least (C. minutilla), and Semipalmated (C. pusilla) sandpipers. We compared the apparent reliability of the isotope approach to that of plumage‐based aging. The large expected differences in δD_f values of flight feathers grown at Arctic versus non‐Arctic latitudes enabled use of this technique to discriminate between age‐classes. We determined δD_f values of known Arctic‐grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather δD_f values of southward‐migrating adults showed bimodal distributions for all three species. Negative values overlapped with species‐specific juvenile values, identifying putative second fall birds with high‐latitude grown juvenal feathers retained from the previous year. The more positive values identified older adults who grew their feathers at mid‐ and low latitudes. Importantly, δD_f analysis successfully identified first‐winter and second‐fall birds not detected by plumage‐based aging. Flight feather wear alone was a poor basis for age classification because scores overlapped extensively between putative second fall birds and older adults. Flight feather hydrogen isotope analysis enables more definitive assignment of age classes when standard plumage methods are unreliable.     

Of 11 candidate steroids,corticosterone concentration standardized for mass is the most reliable steroid biomarker of nutritional stress across different feather types

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