Search

Once a year, penguins undergo a catastrophic moult, replacing their entire plumage during a fasting period on land or on sea-ice during which time individuals can lose 45% of their body mass. In penguins, new feather synthesis precedes the loss of old feathers, leading to an accumulation of two feather layers (double coat) before the old plumage is shed. We hypothesized that the combination of the high metabolism required for new feather synthesis and the potentially high thermal insulation linked to the double coat could lead to a thermal challenge requiring additional peripheral circulation to thermal windows to dissipate the extra heat. To test this hypothesis, we measured the surface temperature of different body regions of captive gentoo penguins (Pygoscelis papua) throughout the moult under constant environmental conditions. The surface temperature of the main body trunk decreased during the initial stages of the moult, suggesting greater thermal insulation. In contrast, the periorbital region, a potential proxy of core temperature in birds, increased during these same early moulting stages. The surface temperature of the bill, flipper and foot (thermal windows) tended to initially increase during the moult, highlighting the likely need for extra heat dissipation in moulting penguins. These results raise questions regarding the thermoregulatory capacities of penguins in the wild during the challenging period of moulting on land in the current context of global warming., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

The evolutionary history of many organisms is characterized by major changes in morphology and distribution. Specifically, alterations of body mass and geographic distribution may profoundly influence organismal life-history traits. Here, we reconstructed the evolutionary history of flight-feather molt strategy using data from 1,808 Neornithes species. Our analysis suggests that the ancestral molt strategy of first-year birds was partial or entirely absent, and that complete wing flight-feather molt in first-year birds first evolved in the late Eocene and Oligocene (25-40 Ma), at least 30 Myr after birds first evolved. Complete flight-feather molt occurred mainly at equatorial latitudes and in relatively low body mass species, following a diversification of body mass within the lineage. We conclude that both body mass and geographic distribution shaped the evolution of molt strategies and propose that the evolutionary transition towards complete juvenile molt in the Neornithes is a novel, relatively late adaptation., (© 2021. The Author(s).)

How dermis maintains tissue homeostasis in cyclic growth and wounding is a fundamental unsolved question. Here, we study how dermal components of feather follicles undergo physiological (molting) and plucking injury-induced regeneration in chickens. Proliferation analyses reveal quiescent, transient-amplifying (TA) and long-term label-retaining dermal cell (LRDC) states. During the growth phase, LRDCs are activated to make new dermal components with distinct cellular flows. Dermal TA cells, enriched in the proximal follicle, generate both peripheral pulp, which extends distally to expand the epithelial-mesenchymal interactive interface for barb patterning, and central pulp, which provides nutrition. Entering the resting phase, LRDCs, accompanying collar bulge epidermal label-retaining cells, descend to the apical dermal papilla. In the next cycle, these apical dermal papilla LRDCs are re-activated to become new pulp progenitor TA cells. In the growth phase, lower dermal sheath can generate dermal papilla and pulp. Transcriptome analyses identify marker genes and highlight molecular signaling associated with dermal specification. We compare the cyclic topological changes with those of the hair follicle, a convergently evolved follicle configuration. This work presents a model for analyzing homeostasis and tissue remodeling of mesenchymal progenitors., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Oxygen, hydrogen, carbon and nitrogen stable isotopes were measured on a comprehensive sampling of feathers from two spring Hooded Warblers (Setophaga citrina) in Texas to evaluate isotopic variability between feathers and during molt. Isotopic homogeneity within each bird was found across all four isotopic systems, supporting the hypothesis that molt in these neotropical migrants is fully completed on the breeding grounds. This homogeneity suggests that the isotopic composition of a single feather is may be representative of the whole songbird. However, each bird was found to have one or two outlier feathers, which could signify regrowth of lost feathers after prebasic molt., Competing Interests: The authors have declared that no competing interests exist.

Modern flying birds molt to replace old and worn feathers that inhibit flight performance, but its origins are unclear. We address this by presenting and evaluating a ~150 million year old record of molting in a feathered dinosaur from the early bird Archaeopteryx. Laser-Stimulated Fluorescence revealed feather sheaths that are otherwise invisible under white light. These are separated by one feather and are not in numerical sequential order and are mirrored in both wings. This indicates that a sequential center-out molting strategy was already present at the origins of flight, which is used in living falcons to preserve maximum flight performance. This strategy would have been a welcome advantage for early theropod flyers that had poor flight capabilities. This discovery provides important insights into how birds refined their early flight capabilities before the appearance of the keeled sternum, pygostyle and triosseal canal.

Feather molt is an important life-history process in birds, but little is known about its evolutionary history. Here, we report on the first fossilized evidence of sequential wing feather molt, a common strategy among extant birds, identified in the Early Cretaceous four-winged dromaeosaurid Microraptor. Analysis of wing feather molt patterns and ecological properties in extant birds imply that Microraptor maintained its flight ability throughout the entire annual cycle, including the molt period. Therefore, we conclude that flight was essential for either its daily foraging or escaping from predators. Our findings propose that the development of sequential molt is the outcome of evolutionary forces to maintain flight capability throughout the entire annual cycle in both extant birds and non-avialan paravian dinosaurs from 120 mya. VIDEO ABSTRACT., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Introduced bird species can become invasive in agroecosystems and their management is inhibited if their origin and movements are not well understood. Stable isotope measurements of feathers can be used to infer molt origins and interstate movements in North America. We analyzed stable-hydrogen (δ2H), carbon (δ13C) and nitrogen (δ 15N) isotope ratios in feathers to better understand the molt origin of European starlings (Sturnus vulgaris) collected at dairies and feedlots throughout the United States. Primary feathers were used from 596 adult and 90 juvenile starlings collected during winter at dairies and feedlots that experience starling damages in Arizona, California, Colorado, Idaho, Illinois, Iowa, Kansas, Minnesota, Missouri, Nebraska, Nevada, New Hampshire, New Mexico, New York, North Carolina, Oregon, Texas, Vermont, Washington and Wisconsin. The best-fit model indicated that the combination of feather δ2H, δ13C and δ15N values best predicted the state where samples were collected and thus supported use of this approach for tracing molt origins in European starlings. Interestingly, molt origins of starlings collected at dairies and feedlots generally west of -90° longitude (i.e. 11 of 15 states west of the Mississippi River, including Wisconsin) were assigned to the collection state and/or the state adjacent to the collection state. In contrast, molt origins of starlings collected generally east of -90° longitude (four of five eastern states) were not assigned to the collection state and/or the state adjacent to the collection state. Among all starlings (N = 686), 23% were assigned to the collection state and 19% were assigned to the state adjacent to the collection state. Among all males (N = 489) and all females (N = 197), 23% and 26% were assigned to the collection state and 19% and 13% were assigned to the state adjacent to the collection state, respectively. We observed a greater proportion (88%) of juvenile starlings assigned to states other than their collection state (i.e. potentially a result of natal dispersal) than that proportion (76%) in adult starlings. This study included an unprecedented sample of feather isotopes from European starlings throughout the United States. As a novel contribution to the ecology and management of invasive and migratory passerines, we demonstrate how such feather isoscapes can be used to predict molt origin and, potentially, interstate movements of European starlings for subsequent ecological and management investigations., Competing Interests: The authors have declared that no competing interests exist.

Abnormal molting, including partial or incomplete molt, arrested molt cycle, or inappropriate frequency of molt, is a primary concern for the managed African penguin ( Spheniscus demersus ) population and is documented across institutions. To identify factors associated with increased odds of abnormal molts and characterize intervention opportunities, a comprehensive survey evaluating numerous husbandry and medical parameters was created. Survey results represent 45 North American African penguin holding facilities and 736 unique animals. Of these individuals, 135 (18.3%) demonstrated an abnormal molt over the 5-yr study period (2012-2017). Increased odds ratios for abnormal molt included biologic (age, sex, etc.), geographic (elevation, latitude), and husbandry (exhibit design, diet, etc.) variables. The mean age of affected animals was 15.2 yr (1-45 yr, n = 135) compared with 9.92 yr (4 mo-38 yr, n = 601) for unaffected animals. In addition, although statistically insignificant, males were overrepresented in the affected cohort compared with a near even distribution among unaffected animals. Identified factors with increased odds for abnormal molting included advanced age and facilities using freshwater pools. Normally molting penguins were more commonly found with saltwater pool access and natural lighting exposure. Anecdotal medical intervention attempts are discussed, although further research is needed to define their use. Of attempted interventions, subcutaneous 5.4-mg melatonin implants placed in anticipation of environmental molting cues showed the most promise at inducing catastrophic molt, with 14 of 17 (82.3%) of affected individuals molting normally following this treatment. Survey analysis indicated that abnormal molt is a complex, multifactorial process, and modifiable factors that may predispose animals to abnormally molt exist. Addressing these factors in future exhibit design may mitigate the prevalence of this condition. Despite these efforts, it is likely that medical interventions will be required to aid in the treatment of abnormal molting in this species.

The measurement of corticosterone levels in feathers (fCort) is gaining recognition as an effective means for describing links between stages of the annual cycle in birds. Many seabirds are especially good models for exploring these links, or carryover effects, due to their migratory behavior and reproductive investment in a single-egg clutch. Here, we measure fCort in Cassin's auklet (Ptychoramphus aleuticus) breast feathers at two colonies in British Columbia during 2011, a year of favorable oceanographic conditions, and examine its relationship with egg size. These feathers are grown at sea during the late winter period, 1-2 months prior to egg laying. Assuming that fCort provides some measure of nutritional stress and hypothalamic-pituitary-adrenal axis activity during feather growth, we predicted that fCort would be positively correlated with egg size via increased support for foraging activity during the nutritionally demanding molt stage. We also analyzed the δ 13 C and δ 15 N stable isotope content of breast feathers, measures commonly used to characterize seabird diet composition. Contrary to prediction, neither fCort nor stable isotope ratios were good predictors of egg size. Our results appear to conflict with two previous studies on alcids in which fCort and stable isotopes showed clear links with egg size; however, both studies were conducted in years when oceanographic and foraging conditions were poor. Under these conditions, upregulation of corticosterone may be needed to mediate the energetic tradeoffs between self-maintenance and reproduction, supporting increased foraging effort and thus increasing both the likelihood of reproduction and large egg size. However, when foraging conditions are favorable, we suggest that such tradeoffs and associated physiological constraints are minimized and regulation of egg size may be effectively independent of circulating corticosterone levels and diet type., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The molting cycle of feathers includes an anagen (growth) stage, a likely catagen stage where the feather follicles degenerate, and a resting stage where fully grown feathers remain in their follicles and are functional before molting. However, the cytological changes involved in the resting and molting stages are poorly known, so the results of an ultrastructural analysis of these processes in adult chick feathers are presented here. The study showed that the dermal papilla shrinks, and numerous cells present increased heterochromatin and free collagen fibrils in the extracellular matrix. Degeneration of the germinal epithelium of the follicle-the papillary collar-occurs with an initial substantial contraction of cells followed by an increase in heterochromatin, vesicle and lipid accumulation, and membrane and organelle degeneration. Desmosomes are still present between degenerating epithelial cells, but ribosomes and tonofilaments disappear. This suggests that cell necrosis initially proceeds as a major contraction resembling apoptosis-a process termed necroptosis, which was previously also shown to occur during the formation of barbs and barbules in mature down and pennaceous feathers. This study suggests that, aside from apoptosis, the collar epithelium degenerates due to external factors, in particular the retraction of blood vessels supplying the dermal papilla. In contrast, revascularization of the dermal papilla triggers a new phase of feather growth (anagen).

Fault bars are narrow malformations in feathers oriented almost perpendicular to the rachis where the feather vein and even the rachis may break. Breaks in the barbs and barbules result in small pieces of the feather vein being lost, while breaks in the rachis result in loss of the distal portion of the feather. Here, we provide a comprehensive review of 74 papers on fault bar formation in hopes of providing a clearer approach to their study. First, we review the evidence that the propensity to develop fault bars is modified by natural selection. Given that fault bars persist in the face of survival costs, we conclude that they must be an unfortunate consequence of some alternative adaptation that outweighs the fitness costs of fault bars. Second, we summarize evidence that the development of fault bars is triggered by psychological stress such as that of handling or predation attempts, and that they persist because the sudden contractions of the muscles in the feather follicle that control fright moults also causes the development of fault bars in growing feathers. Third, we review external and physiological (e.g. corticosterone) agents that may affect the likelihood that an acute stress will result in a growing feather exhibiting a fault bar. These modifying factors have often been treated as fundamental causes in the earlier literature on fault bars. Fourth, we then use this classification to propose a tentative model where fault bars of different severity (from light to severe) are the outcome of the interaction between the propensity to produce fault bars (which differs between species, individuals and feather follicles within individuals) and the intensity of the perturbation. This model helps to explain contradictory results in the literature, to identify gaps in our knowledge, and to suggest further studies. Lastly, we discuss ways in which better understanding of fault bars can inform us about other aspects of avian evolutionary ecology, such as the physiology of moult, the integration of moult into avian life cycles, and the strategies used to minimize stress during moult. Moreover, the study of fault bars may be relevant to understanding the aerodynamics of flight and the early evolution of flight., (© 2016 Cambridge Philosophical Society.)

In order to find out the optimum level of crude protein (CP) in the diet of captive Lady Amherst's pheasants (LAP) on molt, 18 male birds were randomly distributed into three groups of six each in an experiment based on completely randomized block design. The CP content of the diets of birds in groups I, II, and III was 13.4, 16.5, and 19.1%, respectively. Intake and apparent balance of nitrogen increased linearly (P < 0.001) as CP content of the diet increased. Intake and utilization of energy, calcium, and phosphorous were similar among groups. Body mass change and growth rate of feathers were significantly (P < 0.01) lower in group I as compared to groups II and III. There was a positive co-relationship between ME intake and change in body weight (R(2) = 0.89, F = 126.4, P < 0.001). Regression analysis indicates that LAP can maintain body mass when ME supply is 122.2 Kcal/kg BW(0.75)/d. Linear relationships between intake and apparent retention of N, Ca, and P as expressed on mg/kg BW(0.75)/d were all significant. Apparent nitrogen retention, and mean feather growth rate was lower in birds fed diet containing 13.4% CP. Feeding of the diets containing 16.5% CP resulted in improved retention of nitrogen, and mean feather growth rate. Further increase in dietary concentration of CP to 19.1% showed no further improvement. It was concluded that a diet containing 16.5% CP would be optimum for Lady Amherst's pheasants during molt., (© 2015 Wiley Periodicals, Inc.)

For birds, unpredictable environments during the energetically stressful times of moulting and breeding are expected to have negative fitness effects. Detecting those effects however, might be difficult if individuals modulate their physiology and/or behaviours in ways to minimize short-term fitness costs. Corticosterone in feathers (CORTf) is thought to provide information on total baseline and stress-induced CORT levels at moulting and is an integrated measure of hypothalamic-pituitary-adrenal activity during the time feathers are grown. We predicted that CORTf levels in northern common eider females would relate to subsequent body condition, reproductive success and survival, in a population of eiders nesting in the eastern Canadian Arctic during a capricious period marked by annual avian cholera outbreaks. We collected CORTf data from feathers grown during previous moult in autumn and data on phenology of subsequent reproduction and survival for 242 eider females over 5 years. Using path analyses, we detected a direct relationship between CORTf and arrival date and body condition the following year. CORTf also had negative indirect relationships with both eider reproductive success and survival of eiders during an avian cholera outbreak. This indirect effect was dramatic with a reduction of approximately 30% in subsequent survival of eiders during an avian cholera outbreak when mean CORTf increased by 1 standard deviation. This study highlights the importance of events or processes occurring during moult on subsequent expression of life-history traits and relation to individual fitness, and shows that information from non-destructive sampling of individuals can track carry-over effects across seasons., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

The ecology of the willow warbler in the north of Western Siberia is considered, and the adaptations that enable the spread of this species to the Subarctic are analyzed. It is established that one of the key factors that caused the change in the range of this species is the northward distribution of shrubs and, hence, the biomass of insects (available food items of these birds).

The objective of this study was to evaluate if extraction of broken feathers outside the molting period was an efficient method to induce growth of new flight feathers as part of the rehabilitation process of raptors with damaged plumage. Primaries, secondaries, and rectrices (four of each) were removed under general anesthesia from 10 American kestrels (Falco sparverius) using two different protocols: with and without filling the follicle with bismuth subnitrate (Orbeseal) to prevent it from sealing. Birds were kept in large aviaries under natural daylight and outdoor temperatures. Rate and quality of growth of new feathers were assessed regularly for 4 mo. Results were significantly different between the types of feathers: 100% of rectrices, 58% of secondaries, and 8% of primaries started to grow within the 4 winter months, and 95% of rectrices, 30% of secondaries, and 0% of primaries completed their growth normally within this period. The use of Orbeseal did not improve the outcome. The tail feathers began to grow between the second and third weeks at about 2.7 mm/day, and growth was completed within 7 wk. Rate, starting time, and duration of remigial growth were less predictable and varied widely among individuals. Although the exact impact of these extractions on the natural molt is still unclear, most of the primaries that were plucked during this study failed to regrow normally during the following molt. In conclusion, plucking damaged tail feathers in kestrels can successfully decrease the duration of rehabilitation. However, this procedure would not be recommended for wing feathers.

Physiological trade-offs mediated by limiting energy, resources or time constrain the simultaneous expression of major functions and can lead to the evolution of temporal separation between demanding activities. In birds, plumage renewal is a demanding activity, which accomplishes fundamental functions, such as allowing thermal insulation, aerodynamics and socio-sexual signaling. Feather renewal is a very expensive and disabling process, and molt is often partitioned from breeding and migration. However, trade-offs between feather renewal and breeding have been only sparsely studied. In barn swallows (Hirundo rustica) breeding in Italy and undergoing molt during wintering in sub-Saharan Africa, we studied this trade-off by removing a tail feather from a large sample of individuals and analyzing growth bar width, reflecting feather growth rate, and length of the growing replacement feather in relation to the stage in the breeding cycle at removal and clutch size. Growth bar width of females and length of the growing replacement feather of both sexes were smaller when the original feather had been removed after clutch initiation. Importantly, in females both growth bar width and replacement feather length were negatively predicted by clutch size, and more strongly so for large clutches and when feather removal occurred immediately after clutch completion. Hence, we found strong, coherent evidence for a trade-off between reproduction, and laying effort in particular, and the ability to generate new feathers. These results support the hypothesis that the derived condition of molting during wintering in long-distance migrants is maintained by the costs of overlapping breeding and molt.

In the course of 15 days, the thyroxine and corticosterone level was measured daily in blood serum of molting starlings, Sturnus vulgaris, by use of enzyme immunodetection method. Revealed are three-day rhythm in changes of thyroxine level and four-day one in changes of corticosterone level, both rhythms being synchronized in different birds. A beginning of growth of new oar feathers coincides with maximum thyroxine concentration in blood serum and also demonstrates a three-day period. In free-living passerine birds (Passer montamus, Parus major, Chloris chloris) a three-day rhythm is found in the dynamics of feather replacement. This rhythm is manifested synchronously in the studied species that differ in timing of post-nuptial molting. Established is the synchronous manifestation of three-day rhythm in changes of thyroxine level and feather replacement during molting in different passerine species and different birds of the same species.

A temporal separation of energetically costly life history events like reproduction and maintenance of the integumentary system is thought to be promoted by selection to avoid trade-offs and maximize fitness. It has therefore remained somewhat of a paradox that certain vertebrate species can undergo both events simultaneously. Identifying potential costs of overlapping two demanding life history stages will further our understanding of the selection pressures that shape the temporal regulation of life history events in vertebrates. We studied free-living tropical Slaty brush-finches (Atlapetes schistaceus), in which individuals spontaneously overlap reproduction and moult or undergo both events in separation. To assess possible costs of such an overlap we quantified feather quality and flight performance of individuals in different states. We determined individual's life history state by measuring gonad size and scoring moult stage, and collected a newly grown 7(th) primary wing feather for later analysis of feather quality. Finally, we quantified flight performance for each individual in the wild. Overlapping individuals produced lighter and shorter wing feathers than individuals just moulting, with females decreasing feather quality more strongly during the overlap than males. Moreover, overlapping individuals had a reduced flight speed during escape flights, while their foraging flight speed was unaffected. Despite overlappers being larger and having a smaller wing area, their lower body mass resulted in a similar wing load as in breeders or moulters. Individuals measured repeatedly in different states also showed significant decreases in feather quality and escape flight speed during the overlap. Reduced escape flight speed may represent a major consequence of the overlap by increasing predation risk. Our data document costs to undergoing two life history stages simultaneously, which likely arise from energetic trade-offs. Impairments in individual quality and performance may represent important factors that select for temporal separation of life history stages in other species.

Background: The trade-off between current and residual reproductive values is central to life history theory, although the possible mechanisms underlying this trade-off are largely unknown. The 'molt constraint' hypothesis suggests that molt and plumage functionality are compromised by the preceding breeding event, yet this candidate mechanism remains insufficiently explored., Methodology/principal Findings: The seasonal change in photoperiod was manipulated to accelerate the molt rate. This treatment simulates the case of naturally late-breeding birds. House sparrows Passer domesticus experiencing accelerated molt developed shorter flight feathers with more fault bars and body feathers with supposedly lower insulation capacity (i.e. shorter, smaller, with a higher barbule density and fewer plumulaceous barbs). However, the wing, tail and primary feather lengths were shorter in fast-molting birds if they had an inferior body condition, which has been largely overlooked in previous studies. The rachis width of flight feathers was not affected by the treatment, but it was still condition-dependent., Conclusions/significance: This study shows that sedentary birds might face evolutionary costs because of the molt rate-feather quality conflict. This is the first study to experimentally demonstrate that (1) molt rate affects several aspects of body feathers as well as flight feathers and (2) the costly effects of rapid molt are condition-specific. We conclude that molt rate and its association with feather quality might be a major mediator of life history trade-offs. Our findings also suggest a novel advantage of early breeding, i.e. the facilitation of slower molt and the condition-dependent regulation of feather growth.

Food limitation greatly affects bird breeding performance, but the effect of nutritive stress on molt has barely been investigated outside of laboratory settings. Here we show changes in molting patterns for an entire native Hawaiian bird community at 1650-1900 m elevation on the Island of Hawaii between 1989-1999 and 2000-2006, associated with severe food limitation throughout the year beginning in 2000. Young birds and adults of all species took longer to complete their molt, including months never or rarely used during the 1989-1999 decade. These included the cold winter months and even the early months of the following breeding season. In addition, more adults of most species initiated their molt one to two months earlier, during the breeding season. Suspended molt, indicated by birds temporarily not molting primary flight feathers during the months of peak primary molt, increased in prevalence. Food limitation reached the point where individuals of all species had asymmetric molt, with different primary flight feathers molted on each wing. These multiple changes in molt, unprecedented in birds, had survival consequences. Adult birds captured during January to March, 2000-2004, had lower survival in four of five species with little effect of extended molt. Extended molt may be adaptive for a nutrient stressed bird to survive warm temperatures but not cool winter temperatures that may obliterate the energy savings. The changing molt of Hawaiian birds has many implications for conservation and for understanding life history aspects of molt of tropical birds.

Elevation of glucocorticoid (GC) hormone levels is an integral part of stress response (as well as its termination) and immunomodulation. These hormones are also responsible for mobilizing energy stores by stimulation of gluconeogenesis and inhibition of protein synthesis. Elevation of GCs is thus incompatible with other protein-demanding processes, such as moult. Previous studies have shown that chronic elevation of GC hormones suppresses feather growth. Here, we asked whether similar effect would also occur in the case of acute GC elevation and induction of an inflammatory response by foreign antigen. We performed an experiment on captive wild-caught greenfinches (Carduelis chloris) injecting birds with phytohaemagglutinin (PHA) and dexamethasone (DEX) in a factorial design. To assess the possible somatic impacts of these manipulations, we removed one of the outermost tail feathers before the experiment and measured mass and rachis diameter and length of the replacement feathers grown in captivity. Immunostimulation by PHA reduced rachis length, but did not affect feather mass or rachis diameter. Single injection of a synthetic GC hormone DEX significantly reduced all three parameters of feather size. Altogether, these findings demonstrate the sensitivity of feather growth to manipulation of immune and adrenal functions. Our results corroborate the somatic costs of immune activation and suggest that even a short-term elevation of GC hormones may induce long-term somatic costs with a potential impact on fitness. Our findings also imply that a single injection of DEX, frequently used as a diagnostic tool, can have lasting effects and researchers must consider this when designing experiments., (© 2011 Wiley Periodicals, Inc.)

Trade-offs between self-maintenance processes can affect life-history evolution. Integument replacement and the stress response both promote self-maintenance and affect survival in vertebrates. Relationships between the two processes have been studied most extensively in birds, where hormonal stress suppression is down regulated during molt in seasonal species, suggesting a resource-based trade-off between the two processes. The only species found to differ are the rock dove and Eurasian tree sparrow, at least one of which performs a very slow molt that may reduce resource demands during feather growth, permitting investment in the stress response. To test for the presence of a molt-stress response trade-off, we measured hormonal stress responsiveness during and outside molt in two additional species with extended molts, red crossbills (Loxia curvirostra) and zebra finches (Taeniopygia guttata). We found that both species maintain hormonal stress responsiveness during molt. Further, a comparative analysis of all available species revealed a strong relationship between molt duration and degree of hormonal suppression. Though our results support trade-off hypotheses, these data can also be explained by alternative hypotheses that have not been formally addressed in the literature. We found a strong relationship between stress suppression and seasonality of breeding and evidence suggesting that the degree of suppression may be either locally adaptable or plastic and responsive to local environmental conditions. We hypothesize that environmental unpredictability favors extended molt duration, which in turn allows for maintenance of the hormonal stress response, and discuss implications of a possible trade-off for the evolution of molt schedules.

Background: Many animals display colorful signals in their integument which convey information about the quality of their bearer. Theoretically, these ornaments incur differential production and/or maintenance costs that enforce their honesty. However, the proximate mechanisms of production costs are poorly understood and contentious in cases of non-carotenoid-based plumage ornaments like the melanin-based badge and depigmented white wing-bar in house sparrows Passer domesticus. Costly life-history events are adaptively separated in time, thus, when reproduction is extended, the time available for molt is curtailed and, in turn, molt rate is accelerated., Methodology/principal Findings: We experimentally accelerated the molt rate by shortening the photoperiod in order to test whether this environmental constraint is mirrored in the expression of plumage ornaments. Sparrows which had undergone an accelerated molt developed smaller badges and less bright wing-bars compared to conspecifics that molted at a natural rate being held at natural-like photoperiod. There was no difference in the brightness of the badge or the size of the wing-bar., Conclusions/significance: These results indicate that the time available for molt and thus the rate at which molt occurs may constrain the expression of melanin-based and depigmented plumage advertisements. This mechanism may lead to the evolution of honest signaling if the onset of molt is condition-dependent through the timing of and/or trade-off between breeding and molt.

The functional life span of feathers is always much less than the potential life span of birds, so feathers must be renewed regularly. But feather renewal entails important energetic, time and performance costs that must be integrated into the annual cycle. Across species the time required to replace flight feather increases disproportionately with body size, resulting in complex, multiple waves of feather replacement in the primaries of many large birds. We describe the rules of flight feather replacement for Hemiprocne mystacea, a small, 60 g tree swift from the New Guinea region. This species breeds and molts in all months of the year, and flight feather molt occurs during breeding in some individuals. H. mystacea is one to be the smallest species for which stepwise replacement of the primaries and secondaries has been documented; yet, primary replacement is extremely slow in this aerial forager, requiring more than 300 days if molt is not interrupted. We used growth bands to show that primaries grow at an average rate of 2.86 mm/d. The 10 primaries are a single molt series, while the 11 secondaries and five rectrices are each broken into two molt series. In large birds stepwise replacement of the primaries serves to increase the rate of primary replacement while minimizing gaps in the wing. But stepwise replacement of the wing quills in H. mystacea proceeds so slowly that it may be a consequence of the ontogeny of stepwise molting, rather than an adaptation, because the average number of growing primaries is probably lower than 1.14 feathers per wing.

We used allometric scaling to explain why the regular replacement of the primary flight feathers requires disproportionately more time for large birds. Primary growth rate scales to mass (M) as M(0.171), whereas the summed length of the primaries scales almost twice as fast (M(0.316)). The ratio of length (mm) to rate (mm/day), which would be the time needed to replace all the primaries one by one, increases as the 0.14 power of mass (M(0.316)/M(0.171) = M(0.145)), illustrating why the time required to replace the primaries is so important to life history evolution in large birds. Smaller birds generally replace all their flight feathers annually, but larger birds that fly while renewing their primaries often extend the primary molt over two or more years. Most flying birds exhibit one of three fundamentally different modes of primary replacement, and the size distributions of birds associated with these replacement modes suggest that birds that replace their primaries in a single wave of molt cannot approach the size of the largest flying birds without first transitioning to a more complex mode of primary replacement. Finally, we propose two models that could account for the 1/6 power allometry between feather growth rate and body mass, both based on a length-to-surface relationship that transforms the linear, cylindrical growing region responsible for producing feather tissue into an essentially two-dimensional structure. These allometric relationships offer a general explanation for flight feather replacement requiring disproportionately more time for large birds., Competing Interests: The authors have declared that no competing interests exist.

Growing evidence suggests that structural feather colours honestly reflect individual quality or body condition but, contrary to pigment-based colours, it is not clear what mechanism links condition to reflectance in structural feather colours. We experimentally accelerated the moult speed of a group of blue tits (Cyanistes caeruleus) by exposing them to a rapidly decreasing photoperiod and compared the spectral characteristics of their structural feather colours with those of control birds. Blue tits were sexually dimorphic on the UV/blue crown and on the white cheek feathers. Moult speed, however, dramatically reduced brightness and the saturation only on the UV/blue crown feathers, whereas structural white on the cheek feathers was basically unaffected by moult speed. Given that the time available for moulting is usually confined to the period between the end of the breeding season and migration or wintering, UV/blue colours, but not structural white, may convey long-term information about an individual's performance during the previous breeding season. The trade-off between fast moulting and structural colour expression may represent a previously unrecognized selective advantage for early-breeding birds.

Understanding the causes of variation in feather colour in free-living migratory birds has been challenging owing to our inability to track individuals during the moulting period when colours are acquired. Using stable-hydrogen isotopes to estimate moulting locality, we show that the carotenoid-based yellow-orange colour of American redstart (Setophaga ruticilla) tail feathers sampled on the wintering grounds in Central America and the Caribbean is related to the location where feathers were grown the previous season across North America. Males that moulted at southerly latitudes were more likely to grow yellowish feathers compared with males that moulted more orange-red feathers further north. Independent samples obtained on both the breeding and the wintering grounds showed that red chroma-an index of carotenoid content-was not related to the mean daily feather growth rate, suggesting that condition during moult did not influence feather colour. Thus, our results support the hypothesis that feather colour is influenced by ecological conditions at the locations where the birds moulted. We suggest that these colour signals may be influenced by geographical variation in diet related to the availability of carotenoids.

It is important to know how different organs 'manage' their stem cells. Both hair and feather follicles show robust regenerative powers that episodically renew the epithelial organ. However, the evolution of feathers (from reptiles to birds) and hairs (from reptiles to mammals) are independent events and their follicular structures result from convergent evolution. Because feathers do not have the anatomical equivalent of a hair follicle bulge, we are interested in determining where their stem cells are localized. By applying long-term label retention, transplantation and DiI tracing to map stem cell activities, here we show that feather follicles contain slow-cycling long-term label-retaining cells (LRCs), transient amplifying cells and differentiating keratinocytes. Each population, located in anatomically distinct regions, undergoes dynamic homeostasis during the feather cycle. In the growing follicle, LRCs are enriched in a 'collar bulge' niche. In the moulting follicle, LRCs shift to populate a papillar ectoderm niche near the dermal papilla. On transplantation, LRCs show multipotentiality. In a three-dimensional view, LRCs are configured as a ring that is horizontally placed in radially symmetric feathers but tilted in bilaterally symmetric feathers. The changing topology of stem cell activities may contribute to the construction of complex feather forms.

Corticosterone (CORT) is seasonally modulated in many passerines, with plasma CORT concentrations lowest during the prebasic molt when all feathers are replaced. To explain why, we proposed that the birds downregulate natural CORT release during molt in order to avoid CORT's degradative effects on proteins and its inhibition of protein synthesis. If CORT exerted these effects during molt, it could slow protein deposition during feather production and potentially result in a longer period of degraded flight performance. To test this hypothesis, either empty or CORT-filled silastic implants were inserted into captive European starlings (Sturnus vulgaris) and white-crowned sparrows (Zonotrichia leucophrys) undergoing induced (feather replacement after plucking) and natural molts. We then measured the rate of feather re-growth by regularly measuring the length of primary, secondary, and tail feathers. CORT implanted birds showed a significantly decreased rate of feather growth compared to control animals. Basal CORT concentrations of induced molt and non-molting birds were also compared but no difference was noted. The results suggest a tradeoff; a complete set of new feathers may be more important to the survival of a bird than the ability of CORT to respond maximally to a stressor.

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.

Many bird species show delayed plumage maturation (DPM), retaining sub-adult plumage until after their first breeding season. Most explanations assume that DPM increases fitness over the breeding season. However, unless birds undergo a full moult before breeding, DPM could also be an adaptation to increase survival over the previous winter. The winter adaptation hypothesis has never been tested owing to the difficulty of measuring overwinter survival. We experimentally tested this hypothesis in North Island robins (Petroica longipes) using a closed island population where we could accurately estimate survival. The experiment involved dyeing 41 juveniles to mimic adult males, and comparing their survival with 41 control juveniles treated with the same peroxide base minus the pigment. The population was monitored with a series of resighting surveys, and mark-recapture analysis used to estimate overwinter survival. Survival probability was estimated to be 10% for dyed birds versus 61% for control birds in 2001, and 29% for dyed birds versus 40% for control birds in the winter of 2002, supporting the winter adaptation hypothesis for DPM. Access to suitable habitat is the key factor limiting juvenile survival in this population, and the locations where dyed juveniles were sighted suggest that they were often excluded from suitable areas.

Life-history theory proposes that costs must be associated with reproduction. Many direct costs are incurred during breeding. There is also evidence for indirect costs, incurred after breeding, which decrease survival and future reproductive success. One possible indirect cost identified in birds is that breeding activity in some way compromises plumage quality in the subsequent moult. Here we propose a mechanism by which this could occur. Breeding activity delays the start of moult. Birds that start to moult later also moult more rapidly--an effect of decreasing daylength. Could this result in poorer quality plumage? We kept two groups of male European starlings, Sturnus vulgaris, one on constant long days and the other on decreasing daylengths from the start of moult. Decreasing daylengths reduced the duration of moult from 103 +/- 4 days to 73 +/- 3 days (p < 0.0001). Newly grown primary feathers of birds that moulted fast were slightly shorter, weighed less (p < 0.05) and were more asymmetrical. They had a thinner rachis (p < 0.005), were less hard (p < 0.01) and less rigid (p < 0.05). They were also less resistant to wear so that differences in mass and asymmetry increased with time. There was no difference in Young's modulus. Poorer quality plumage will lead to decreased survival due to decreased flight performance and increased thermoregulatory costs. Thus, reproduction incurs costs through a mechanism that operates after the end of breeding.