Theory of the development of curved barbs and their effects on feather morphology

Feathers exhibit an extraordinary diversity of shapes, which are used by birds to accomplish a diverse set of functions. Pennaceous feathers have a double branched morphology that develops from a tube of epidermis, and variation in branch geometry determines feather shape. Feather development is both complex (i.e., a simple developmental modification can have multiple effects on mature feather shape), and redundant (i.e., different developmental modifications can create the same shape). Due to this, it is not readily apparent how different feather shapes develop. In many feathers, barbs are not straight, but instead curve in toward, or away, from the feather tip. Barb curvature can affect the shape of mature feathers but the development of curved barbs is unknown. Previous research has hypothesized that barb curvature could develop either during the helical growth of barb ridges in the tubular feather germ, or during barb angle expansion as the feather unfurls from the sheath. To better understand the development of curved barbs and their effects on mature feathers we present a theoretical model of curved barb development and test the model with empirical investigations of feathers. We find that curved barbs affect many aspects of feather morphology including vane width, barb length, and barb spacing. In real feathers, curved barbs can develop both during helical barb ridge growth and during barb angle expansion, with most of the observed curvature due to barb angle expansion. Our results demonstrate that barb angle expansion as a feather unfurls from the sheath is a complex and dynamic process that plays an important role in determining the shape and structure of mature feathers. Curved barbs create heterogeneity in barb geometry within the feather vane, which could have important implications for aerodynamic function and the development of within feather pigmentation patterns. J. Morphol. 277:995–1013, 2016. © 2016 Wiley Periodicals, Inc.     

Feather eating and its associations with plumage damage and feathers on the floor in commercial farms of laying hens

Feather eating has been associated with feather pecking, which continues to pose economic and welfare problems in egg production. Knowledge on feather eating is limited and studies of feather eating in commercial flocks of laying hens have not been performed previously. Therefore, the main objective was to investigate feather eating and its association with plumage damage and floor feather characteristics in commercial flocks of layers in barn and organic production systems. The study was performed in 13 flocks of barn layers and 17 flocks of organic layers. Each flock was visited at around 32 and 62 weeks of age. During both visits, the plumage condition was assessed and the density of floor feathers recorded. In week 62, droppings and floor feathers were collected. Droppings were examined for presence of feather content, whereas length, downiness and pecking damage were recorded for each floor feather. In week 62, a higher prevalence of hens with poor plumage condition was found in barn (22.2%) compared with organic production systems (7.4%; P<0.001), but the prevalence of droppings with feather content did not differ between the two production systems (8.5% in barn v. 4.3% in organic; P=0.99). Our hypothesis about a positive correlation between feather eating and plumage damage was not supported as no correlation was found between the prevalence of poor plumage condition and the prevalence of droppings with feather content. However, the prevalence of pecking damaged floor feathers was positively correlated both with prevalence of droppings with feather content (P<0.05) and poor plumage condition (P<0.01), indicating a possible association between feather eating and feather pecking. In conclusion, it was confirmed that feather eating occurs on-farm, but feather eating was only found to be positively correlated to the number of floor feathers with pecking damage and not as expected to the prevalence of plumage damage. More research is needed into the sources from where feathers are selected for ingestion, that is, whether they are picked from the floor litter, plucked directly from other hens or dislodged during preening of own feathers.     

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.     

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.     

Feather degradation by Kocuria rosea in submerged culture

A strain of Kocuria rosea with keratinolytic activity was studied. In batch culture, the optimum temperature for feather degradation, bacterial growth and protease secretion was at 40 °C. A specific growth rate of 0.17 h⁻¹ was attained in basal medium with feathers as fermentation substrate. Under these conditions, after 36 h of incubation, biomass and caseinolytic activity reached 3.2 g/l and 0.15 U/ml, respectively. Extracellular protease secretion was associated with the exponential growth phase. In batch fermentation, feather degradation up to 51% in 72 h was obtained with a conversion yield in biomass of 0.32 g/g. No organic acids were detected in the fermentation broth in significant amount. This revised version was published online in July 2006 with corrections to the Cover Date.     

Web databases of feather photographs are useful tools for avian morphometry studies

1. Wing area, wing loading, and aspect ratio are key variables for studies of avian comparative ecology, despite the complexity of measuring wing characteristics in living and museum specimens. The systematic databases of feather photographs available on the Internet may offer an alternative way of obtaining such morphometric data. Here, we evaluate whether measurements of scanned feathers from web photograph databases may offer reliable estimates of avian morphometry.
2. Published data on wing area were obtained for 317 bird species and feather measurements from web photograph databases for 225 of them. A variable termed “lift generation area,” a proxy for wing area, was calculated for each species on the basis of the mean length of the five distal secondary feathers and wingspan data from literature. The fit between this proposed variable and data extracted from the literature was examined by correlation, employing linear regression to explore the lack of fit among species.
3. “Lift generation area” proved to be highly informative as a proxy for wing area for the study species as a whole (R ² > .98). Discrepancies observed between species were strongly negatively associated with the size of the original sample used to calculate wing area (p = .001) and, to a lesser extent, with bird size (p = .023), but not with aspect ratio. It was also found that the mean value of the mismatch between “lift generation area” and wing area (13.1%) among the study species as a whole was of similar magnitude to that found between sources of bibliographic wing area data for the 64 species for which two published estimates of this variable were available (15.3%).
4. We conclude that measurements made from feather photograph databases are reliable for use in studies of avian comparative ecology, enabling the inclusion of biomechanical parameters of many more species than featured at present.

     

House Sparrows Passer domesticus with larger uropygial glands show reduced feather wear

This study assesses whether uropygial gland size is related to improved feather quality. To address this question, I analysed the relationship between uropygial gland size and feather wear in the House Sparrow Passer domesticus. The results show that birds with larger uropygial glands had less worn feathers, suggesting that uropygial gland secretions improve feather resistance to abrasion.     

Growth and annual survival estimates to examine the ecology of larval lamprey and the implications of ageing error in fitting models

This study used existing western brook lamprey Lampetra richardsoni age information to fit three different growth models (i.e. von Bertalanffy, Gompertz and logistic) with and without error in age estimates. Among these growth models, there was greater support for the logistic and Gompertz models than the von Bertalanffy model, regardless of ageing error assumptions. The von Bertalanffy model, however, appeared to fit the data well enough to permit survival estimates; using length‐based estimators, annual survival varied between 0·64 (95% credibility interval: 0·44–0·79) and 0·81 (0·79–0·83) depending on ageing and growth process error structure. These estimates are applicable to conservation and management of L. richardsoni and other western lampreys (e.g. Pacific lamprey Entosphenus tridentatus) and can potentially be used in the development of life‐cycle models for these species. These results also suggest that estimators derived from von Bertalanffy growth models should be interpreted with caution if there is high uncertainty in age estimates.     

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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Parasites and the blackcap's tail: implications for the evolution of feather ornaments

Although parasites may impair the expression of tail ornaments in birds, the importance of parasitism in driving the evolution of the initial stages of tail ornamentation is not well understood. Parasites could have negatively affected the expression of nonexaggerated, functional traits before these evolved ornaments, or they could have played a relevant role only after tails became ornamental and hence too costly to produce. To shed light on this issue, we studied the correlation between the abundance of feather mites (Acari, Proctophyllodidae) and the size, quality, growth rate and symmetry of tail feathers of blackcaps ( Sylvia atricapilla ), a non-ornamented passerine. Tail length was not correlated with mite load, yet blackcaps holding many mites at the moment of feather growth (fledglings) had lighter and more asymmetric feathers that grew at relatively lower rates. In blackcaps whose mite load was measured one year after feather growth (adults), only the negative correlation between mite intensity and feather symmetry remained significant. Changes in mite load since the moult season could have erased the correlation between condition-dependent feather traits and current parasite load in adults. Our results support the idea that different traits of non-ornamental feathers can signal parasite resistance. Therefore, parasitism could have played a central role in the evolution of tail ornamentation ever since its initial stages.  © 2002 The Linnean Society of London. Biological Journal of the Linnean Society , 2002, 76 , 481–492.     

Characterization of a new keratinolytic Trichoderma atroviride strain F6 that completely degrades native chicken feather

Aims:  To isolate novel nonpathogenic fungus that completely degrades native chicken feather and characterize its keratinases. Methods and Results:  Feather‐degrading fungi were isolated from decaying feathers using a novel method based on simulating decaying process in the environment. The isolate F6 with high keratinolytic activity was identified as Trichoderma atroviride based on morphological traits and ITS1‐5·8S‐ITS2 sequence analysis. The purified dominant component of keratinase had a molecular mass of 21 kDa. The purified keratinase belonged to serine protease. Its isoelectric point, molecular weight, optimum pH, optimum temperature, and substrate specificity are different from those of other serine proteases of Trichoderma species. The optimum pH and temperature values of purified keratinase were consistent with those of crude keratinase. However, the differences between crude and purified enzymes such as thermostability, resistance to Ba²⁺, Mn²⁺, Hg²⁺, Zn²⁺, Cu²⁺, 1,10‐phenanthroline, 2,2′‐bipyridyl, and PMSF (phenylmethylsulfonyl fluoride) were observed. Conclusions:  The results suggested the purified keratinase is predominantly extracellular proteins when strain F6 was grown on keratinous substrates. The protease, in combination with other components, is effective in feather degradation. The strain F6 is more suitable for feather degradation than its purified keratinase. Significance and Impact of the Study:  The novel nonpathogenic T. atroviride F6 with high feather‐degrading activity showed potentials in biotechnological process of converting feathers into economically useful feather meal.     

Barb geometry of asymmetrical feathers reveals a transitional morphology in the evolution of avian flight

The geometry of feather barbs (barb length and barb angle) determines feather vane asymmetry and vane rigidity, which are both critical to a feather''s aerodynamic performance. Here, we describe the relationship between barb geometry and aerodynamic function across the evolutionary history of asymmetrical flight feathers, from Mesozoic taxa outside of modern avian diversity (Microraptor, Archaeopteryx, Sapeornis, Confuciusornis and the enantiornithine Eopengornis) to an extensive sample of modern birds. Contrary to previous assumptions, we find that barb angle is not related to vane-width asymmetry; instead barb angle varies with vane function, whereas barb length variation determines vane asymmetry. We demonstrate that barb geometry significantly differs among functionally distinct portions of flight feather vanes, and that cutting-edge leading vanes occupy a distinct region of morphospace characterized by small barb angles. This cutting-edge vane morphology is ubiquitous across a phylogenetically and functionally diverse sample of modern birds and Mesozoic stem birds, revealing a fundamental aerodynamic adaptation that has persisted from the Late Jurassic. However, in Mesozoic taxa stemward of Ornithurae and Enantiornithes, trailing vane barb geometry is distinctly different from that of modern birds. In both modern birds and enantiornithines, trailing vanes have larger barb angles than in comparatively stemward taxa like Archaeopteryx, which exhibit small trailing vane barb angles. This discovery reveals a previously unrecognized evolutionary transition in flight feather morphology, which has important implications for the flight capacity of early feathered theropods such as Archaeopteryx and Microraptor. Our findings suggest that the fully modern avian flight feather, and possibly a modern capacity for powered flight, evolved crownward of Confuciusornis, long after the origin of asymmetrical flight feathers, and much later than previously recognized.     

[树]麻雀羽再生的能量预算和水代谢散热调节

为探讨[树]麻雀的羽再生能力、能量预算对策和有效的散热调节方式,对3组(对照CF、去飞羽FF和去尾羽组TF)[树]麻雀(Passer montanus)进行4周驯养(Acclimation)。结果发现:[树]麻雀具有较强的羽再生能力和飞羽参与个体保温。羽再生[树]麻雀能量收支水平降低极显著(P﹤0.001),FF组和TF组比CF组减少依次为:摄入能19.77%和7.17%、消化能18.79%和6.47%、同化能18.73%和6.46%、粪能28.66%和13.35%、水代谢散热热能26.95%和7.43%、排泄次数33.71%和14.40%,增加依次为:消化率1.23%和0.78%、同化率1.35%和0.84%。个体能值水平,体重CF、TF和FF组(P﹤0.05)依次降低,体温组间变化不显著(P﹥0.05)。体内能量储备,血糖、肌糖原、体脂和水分含量组间差异不显著(P﹥0.05),肝糖原含量、体脂重组间差异显著(P﹤0.05)。器官水平包括心脏、肾脏、腺胃、小肠、盲肠和总消化道长度及质量出现积极的响应。日代谢水量组间差异极显著(P﹤0.001)。组间日排泄次数最少平均为56.11次和最多可达96.34次/只。结果表明:羽再生[树]麻雀分别选择了不同程度的降低能量收支水平,提高摄入食物的消化、吸收效率,动用体内能量储备来获取摄入能量不足部分,降低器官总能耗的能量预算对策和不同的新羽(再生羽枚数:飞羽部分和尾羽全部)再生的能量投资对策。泄殖腔排出(粪尿混合物)水是鸟类特有的、迅速的和有效的散热调节方式。     

Comparative analysis of nonlinear growth curve models for <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> rosette leaves

As a model organism, modeling and analysis of the phenotype of Arabidopsis thaliana (A. thaliana) leaves for a given genotype can help us better understand leaf growth regulation. A. thaliana leaves growth trajectories are to be nonlinear and the leaves contribute most to the above-ground biomass. Therefore, analysis of their change regulation and development of nonlinear growth models can better understand the phenotypic characteristics of leaves (e.g., leaf size) at different growth stages. In this study, every individual leaf size of A. thaliana rosette leaves was measured during their whole life cycle using non-destructive imaging measurement. And three growth models (Gompertz model, logistic model and Von Bertalanffy model) were analyzed to quantify the rosette leaves growth process of A. thaliana. Both graphical (plots of standardized residuals) and numerical measures (AIC, R² and RMSE) were used to evaluate the fitted models. The results showed that the logistic model fitted better in describing the growth of A. thaliana leaves compared to Gompertz model and Von Bertalanffy model, as it gave higher R² and lower AIC and RMSE for the leaves of A. thaliana at different growth stages (i.e., early leaf, mid-term leaf and late leaf).     

Toward a Deuterium Feather Isoscape for Sub-Saharan Africa: Progress,Challenges and the Path Ahead

A key challenge to the application of continent-wide feather isoscapes for geographic assignment of migrant birds is the lack of ground-truthed samples. This is especially true for long-distance Palearctic-Afrotropical migrants. We used spatially-explicit information on the δ ²H composition of archived feathers from Green-backed/Grey-backed Camaroptera, to create a feather δ ²H isoscape for sub-Saharan Africa. We sampled from 34 out of 41 sub-Saharan countries, totaling 205 sampling localities. Feather samples were obtained from museum collections (n = 224, from 1950 to 2014) for δ ²H assay. Region, altitude, annual rainfall and seasonal patterns in precipitation were revealed as relevant explanatory variables for spatial patterns in feather δ ²H. Predicted feather δ ²H values ranged from -4.0 ‰ to -63.3 ‰, with higher values observed in the Great Rift Valley and South Africa, and lower values in central Africa. Our feather isoscape differed from that modelled previously using a precipitation δ ²H isoscape and an assumed feather-to-precipitation calibration, but the relatively low model goodness fit (F_10,213 = 5.98, p<0.001, R² = 0.18) suggests that other, non-controlled variables might be driving observed geographic patterns in feather δ ²H values. Additional ground-truthing studies are therefore recommended to improve the accuracy of the African feather δ ²H isoscape.     

Assessment of the effect of housing on feather damage in laying hens using IR thermography

Plumage damage represents one of the animal-based measures of laying hens welfare. Damage occurs predominantly due to age, environment and damaging pecking. IR thermography, due to its non-invasiveness, objectivity and repeatability is a promising alternative to feather damage scoring systems such as the system included in the Welfare Quality^R assessment protocol for poultry. The aim of this study was to apply IR thermography for the assessment of feather damage in laying hens kept in two housing systems and to compare the results with feather scoring. At the start of the experiment, 16-week-old laying hens (n=30) were divided into two treatments such as deep litter pen and enriched cage. During 4 months, feather damage was assessed regularly in 2-week intervals. One more single assessment was done nine and a half months after the start of the experiment. The feather damage on four body regions was assessed by scoring and IR thermography: head and neck, back and rump, belly, and underneck and breast. Two variables obtained by IR thermography were used: the difference between the body surface temperature and ambient temperature (ΔT_B) and the proportion of featherless areas, which were defined as areas with a temperature >33.5°C. Data were analyzed using a GLM model. The effects of housing, time, region and their interactions on feather damage, measured by the feather scoring and by both IR thermography measures, were all significant (P<0.001). The ΔT_B in all assessed regions correlated positively with the feather score. Feather scoring revealed higher damage in enriched cages compared with deep litter pens starting from week 6 of the experiment on the belly and back and rump regions, whereas ΔT_B from week 6 in the belly and from week 8 on the back and rump region. The proportion of featherless areas in the belly region differed significantly between the housings from week 8 of the experiment and on the back and rump region from week 12. The IR thermography assessment of the feather damage revealed differences between hens kept in different housing systems in agreement with the feather scoring. In conclusion, it was demonstrated that the IR thermography is a useful tool for the assessment of poultry feather cover quality that is not biased by the subjective component and provides higher precision than feather damage scoring.     

Adventitious feather replacement favours a more rapid regeneration of primaries over rectrices in two passerine bird species

There is increasing evidence of adaptive preferential investment during moult in those feather tracts that are more advantageous for fitness. In this study, we assessed whether, after the manual removal of two functionally different flight feathers (one primary and one rectrix), birds from two common passerine species (Eurasian Blackcap Sylvia atricapilla and European Robin Erithacus rubecula) favoured the regeneration of primary (supposedly the most functionally important feathers) over rectrix feathers. Our results did not show differences between replaced primary and rectrix feathers in their final length, but demonstrated that the gap left by the loss of the primary feather was filled earlier, suggesting that a rapid repair of the most essential feather tracts is also evolutionarily advantageous during the adventitious replacement of plumage.     

Isolation and characterization of a feather-degrading bacterium from the poultry processing industry

A Flavobacterium sp. producing a high keratinolytic activity was isolated from a poultry industry after growth on selective feather meal agar. This bacterium grew on feather meal broth, producing keratinase, and was also capable of complete degradation of raw feathers. The proteolytic activity was assessed in the presence of specific protease inhibitors. The crude enzyme showed mainly metalloprotease character. This novel isolate would have potential biotechnological use in processes involving keratin hydrolysis. Received 09 October 2001/ Accepted in revised form 19 July 2002     

Age and growth of the sandbar shark, Carcharhinus plumbeus, in Hawaiian waters through vertebral analysis

Age and growth estimates were determined for the sandbar shark, Carcharhinus plumbeus, from Oahu, Hawaii in the central Pacific Ocean. Age estimates were obtained through vertebral centra analysis of 187 sharks. We verified our age estimates through marginal increment analysis of centra and oxytetracycline marking methods of at liberty sandbar sharks. Sizes of sampled sharks ranged from 46 to 147 cm pre-caudal length. Four growth models were fitted to length-at-age data; two forms of the von Bertalanffy growth model, the Gompertz growth model, and a logistic growth model. Males and females exhibited statistically significant differences in growth, indicating that females grow slower and attain larger sizes than males. Growth parameter estimates revealed slower growth rates than previously estimated (based on captive specimens) for Hawaiian sandbar sharks. The von Bertalanffy growth model using empirical length-at-birth provided the best biological and statistical fit to the data. This model gave parameter estimates of L _∞ = 138.5 cm PCL and k = 0.12 year⁻¹ for males and L _∞ = 152.8 cm PCL, k = 0.10 year⁻¹ for females. Male and female sandbar sharks mature at approximately 8 and 10 years of age, respectively.     

Isolation, Identification, and Characterization of a Feather-Degrading Bacterium

A feather-degrading culture was enriched with isolates from a poultry waste digestor and adapted to grow with feathers as its primary source of carbon, sulfur, and energy. Subsequently, a feather-hydrolytic, endospore-forming, motile, rod-shaped bacterium was isolated from the feather-degrading culture. The organism was Gram stain variable and catalase positive and demonstrated facultative growth at thermophilic temperatures. The optimum rate of growth in nutrient broth occurred at 45 to 50°C and at pH 7.5. Electron microscopy of the isolate showed internal crystals. The microorganism was identified as Bacillus licheniformis PWD-1. Growth on hammer-milled-feather medium of various substrate concentrations was determined by plate colony count. Maximum growth (approximately 10⁹ cells per ml) at 50°C occurred 5 days postinoculation on 1% feather substrate. Feather hydrolysis was evidenced as free amino acids produced in the medium. The most efficient conditions for feather fermentation occurred during the incubation of 1 part feathers to 2 parts B. licheniformis PWD-1 culture (10⁷ cells per ml) for 6 days at 50°C. These data indicate a potential biotechnique for degradation and utilization of feather keratin.