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Evolution of the morphological innovations of feathers.

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

Animals↗

Differentiation of embryonic chick feather-forming and scale-forming tissues in transfilter cultures.

The dermal-epidermal tissue interaction in the chick embryo, leading to the formation of feathers and scales, provides a good experimental system to study the transfer between tissues of signals which specify cell type. At certain times in development, the dermis controls whether the epidermis forms feathers or scales, each of which are characterized by the synthesis of specific beta-keratins. In our culture system, a dermal effect on epidermal differentiation can still be observed, even when the tissues are separated by a Nuclepore filter, although development is abnormal. Epidermal morphological and histological differentiation in transfilter cultures are distinct and recognizable, more closely resembling feather or scale development, depending on the regional origin of the dermis. Differentiation is more advanced when epidermis is cultured transfilter from scale dermis than from feather dermis, as assessed by morphology and histology, as well as the expression of the tissue-specific gene products, the beta-keratins. Two-dimensional polyacrylamide gel analysis of the beta-keratins reveals that scale dermis cultured transfilter from either presumptive scale or feather epidermis induces the production of 7 of the 9 scale-specific beta-keratins that we have identified. Feather dermis, although less effective in activating the feather gene program when cultured transfilter from either presumptive feather or scale epidermis, is able to turn on the synthesis of 3 to 6 of the 18 feather-specific beta-keratins that we have identified. However, scale epidermis in transfilter recombinants with feather dermis also continues to synthesize many of the scale-specific beta-keratins. Using transmission and scanning electron microscopy, we detect no cell contact between tissues separated by a 0.2-micron pore diameter Nuclepore filter, while 0.4-micron filters readily permit cell processes to traverse the filter. We find that epidermal differentiation is the same with either pore size filter. Furthermore, we do not detect a basement membrane in transfilter cultures, implying that neither direct cell contact between dermis and epidermis, nor a basement membrane between the tissues is required for the extent of epidermal differentiation that we observe.

Animals↗

A clinical study of hypersensitivity pneumonitis presumably caused by feather duvets.

BACKGROUND: Bird fancier's lung (BFL) is a type of hypersensitivity pneumonitis induced by the inhalation of bird-related antigens. The BFL induced by feathers is difficult to diagnose because feathers are generally unrecognized as a causative antigen. OBJECTIVE: To determine the clinical features of BFL presumably induced by feather duvets (feather duvet lung) to provide clues for diagnosis. METHODS: We performed a retrospective review of the medical records of patients with feather duvet lung evaluated between April 1, 2000, and June 30, 2003, at the Tokyo Medical and Dental University Hospital in Japan. RESULTS: Seven patients with feather duvet lung were included in this study; 4 patients had acute disease and 3 had chronic BFL. Duration of contact with feather duvets was 1 month to 10 years. Serum KL-6 and surfactant protein D levels were elevated in all the patients. Specific antibodies against avian antigens were positive in acute BFL but negative in chronic BFL. Antigen-induced lymphocyte proliferation in peripheral blood or bronchoalveolar lavage cells was positive in all the patients. The diagnosis was confirmed by an environmental or inhalation provocation test. CONCLUSIONS: Feather duvets can induce acute and chronic BFL. Physicians should be aware of feather duvets as a cause of BFL because feather duvets are becoming more prevalent.

Adult↗

Morphometry of auricular feathers of barn owls (Tyto alba).

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

Animals↗

Feather pecking in growers: a study with individually marked birds.

1. The aim of the present study was to investigate whether individual birds specialise in feather pecking. Growers were individually marked and reared in groups of 30 or 31 in pens with a slatted floor. At an age of 4 to 6 weeks feather pecking was frequent in all pens. 2. On average 83% of all group members (10 groups, experiment 1) were recorded at least once as initiator of a feather pecking interaction. In each group 2 to 6 individuals feather pecked more than twice as often as the average for the group, and were defined as 'high rate peckers'. They initiated 39% of all recorded feather pecking interactions. 3. Every interaction was classified (with increasing intensity) as pecking, pinching, pulling or plucking. Compared to the others, 'high rate peckers' had more of their feather pecking classified as plucking and less classified as pecking. 4. There was no evidence that particular individuals specialised in pecking at other specific birds, at specific areas of the body or at birds engaged in specific activities. 5. Growers (3 groups, experiment 2) that had just feather pecked engaged in more feather pecking during a subsequent 2-min focal observation than control birds that had not pecked before the start of the observation. 6. It was concluded that feather pecking interactions are clustered in time and that the behaviour is not performed by just a few members of a group. However, some individuals are characterised by relatively high rates and more severe forms of feather pecking.

Aggression↗

Methionine and cystine requirements of slow- and fast-feathering broiler males from three to six weeks of age.

Two experiments were conducted to first determine Met then Cys needs of broilers from 3 to 6 wk of age and whether differences existed between slow-feathering (Ross x 308) and fast-feathering (Ross x 3F8) males. A corn-soybean meal diet (20.0% CP; 3,150 kcal ME/kg) with graded levels of Met or Cys was offered. The first experiment had dietary Met levels of 0.32, 0.38, 0.44, and 0.50% with surfeit Cys (0.40%). Broilers from both feathering strains responded similarly to supplemental Met. Although body weight was not responsive, F/G improved through to the highest level of dietary Met (linear, P < 0.05). Chilled carcass weight increased with Met (linear, P < 0.05) paralleling F/G; however, no differences were detected in the amount of associated abdominal fat. Breast fillet yield increased with Met to maximize at 0.48% (quadratic, P < or = 0.009). In a satellite study using the same birds in cages and feeds, N retention at d 29 maximized at 0.46% Met (quadratic, P < 0.05). The second experiment had Cys at 0.32, 0.34, 0.38, and 0.46% with Met fixed at a submarginal level of 0.38%. Increasing dietary Cys had no effect on live performance of slow-feathering birds, whereas weight gain of fast-feathering birds achieved maximum at 0.36% Cys (cubic; P < 0.05) with F/G responding similarly. Chilled carcass (cubic, P < 0.002) and breast fillet weights (cubic, P < 0.001) of fast-feathering birds also increased with Cys to maximize at 0.36%, and the amount of abdominal fat was not influenced by feathering or Cys supplementation. Separate measurement of N retention at d 31 failed to detect a difference in protein utilization attributable to feathering, but an optimum was achieved at 0.40% Cys with both broiler sources. Overall results suggest that the Met requirement for broiler males between 3 and 6 wk of age was independent of feathering and approximated 0.46% (95% of the level of maximal response). Cystine requirements once corrected for submarginal Met status indicated a greater demand by fast- than slow-feathering male broilers corresponding to 0.42 and 0.37%, respectively.

Age Factors↗

FGF induces new feather buds from developing avian skin.

Induction of skin appendages involves a cascade of molecular events. The fibroblast growth factor (FGF) family of peptide growth factors is involved in cell proliferation and morphogenesis. We explored the role of the FGFs during skin appendage induction using developing chicken feather buds as a model. FGF-1, FGF-2, or FGF-4 was added directly to the culture medium or was released from pre-soaked Affigel blue beads. Near the midline, FGFs led to fusion of developing feather buds, representing FGFs' ability to expand feather bud domains in developing skin. In lateral regions of the explant where feather placodes have not formed, FGF treatment produces a zone of condensation and a region with an increased number of feather buds. In ventral epidermis that is normally apteric (without feathers), FGFs can also induce new feather buds. Like normal feather buds, the newly induced buds express Shh. The expression of Grb, Ras, Raf, and Erk, intracellular signaling molecules known to be downstream to tyrosine kinase receptors such as the FGF receptor, was enriched in feather bud domains. Genistein, an inhibitor of tyrosine kinase, suppressed feather bud formation and the effect of FGF. These results indicate that there are varied responses to FGFs depending on epithelial competence. All the phenotypic responses, however, show that FGFs facilitate the formation of skin appendage domains.

Animals↗

Evolution of the structure of tail feathers: implications for the theory of sexual selection.

Bird tails are extraordinarily variable in length and functionality. In some species, males have evolved exaggeratedly long tails as a result of sexual selection. Changes in tail length should be associated with changes in feather structure. The study of the evolution of feather structure in bird tails could give insight to understand the causes and means of evolution in relation to processes of sexual selection. In theory, three possible means of tail length evolution in relation to structural components might be expected: (1) a positive relationship between the increase in length and size of structural components maintaining the mechanical properties of the feather; (2) no relationship; that is, enlarging feather length without changes in the structural components; and (3) a negative relationship; that is, enlarging feather length by reducing structural components. These hypotheses were tested using phylogenetic analyses to examine changes in both degree of exaggeration in tail length and structural characteristics of tail feathers (rachis width and density of barbs) in 36 species, including those dimorphic and nondimorphic in tail length. The degree of sexual dimorphism in tail length was negatively correlated with both rachis width and density of barbs in males but not in females. Reinforcing this result, we found that dimorphism in tail length was negatively associated with dimorphism in tail feather structure (rachis width and density of barbs). These results support the third hypothesis, in which the evolution of long feathers occurs at the expense of making them simpler and therefore less costly to produce. However, we do not know the effects of enfeeblement on the costs of bearing. If the total costs increased, the enfeeblement of feathers could be explained as a reinforcement of the honesty of the signal. Alternatively, if total costs were reduced, the strategy could be explained by cheating processes. The study of female preferences for fragile tail feathers is essential to test these two hypotheses. Preferences for fragile tails would support the evolution of reinforcement of honesty, whereas female indifference would indicate the existence of cheating in certain stages of the evolutionary process.

Animals↗

Are genetic differences in foraging behaviour of laying hen chicks paralleled by hybrid-specific differences in feather pecking?

Feather pecking is a behavioural disorder in laying hens which consists of pecking the feathers of conspecifics, causing feather damage or even injuries to the skin. Its development can be explained by redirection of foraging behaviour. While the occurrence of feather pecking strongly depends on the kind of housing condition, it is also known that there are strain differences in the tendency to feather peck. From the inverse relation between feather pecking and foraging behaviour found earlier, we hypothesised that genetically determined differences in foraging behaviour could be responsible for the observed differences in feather pecking between strains.In a first experiment we tested whether there are differences in the foraging behaviour of two hybrids. As hybrids, we used Lohman selected leghorn (LSL) and Dekalb; eight groups of 20 1-day old chicks each. They were kept in enriched pens (265cmx90cm) with a litter area (200cmx90cm) consisting of wood-shavings, chaff, straw, polystyrene blocks, sand area (65cmx90cm) and elevated perches. Behavioural observations were carried out in week 4. In a subsequent experiment with the same birds we tested how the foraging behaviour of the two hybrids differed when housing conditions were changed from enriched to restricted and to what extent they developed feather pecking. A 2x2 factorial design with hybrid (LSL, Dekalb) and housing condition (restricted, enriched) as factors and with four replicates of each factor combination was used. Half of the pens of each hybrid were changed from enriched to restricted housing conditions by covering the litter area with slats. Behavioural observations were carried out in weeks 5 and 6.In experiment 1, LSL and Dekalb spent the same amount of time foraging, but Dekalb spent significantly more of that time with pecking and hacking at the polystyrene blocks. On the other hand, LSL spent significantly more time at the feeding troughs and rested significantly less than Dekalb. In the restricted environment of experiment 2, again, the total foraging time did not differ between hybrids, but LSL chicks spent significantly less time scratching, while Dekalb spent significantly more time moving. Both hybrids developed feather pecking but LSL showed significantly higher rates than Dekalb.Our results demonstrate genetic differences in the foraging behaviour and in the way hybrids cope with the change in housing condition from enriched to an environment that is restricted in relation to foraging possibilities. We conclude that the results support the hypothesis put forward that genetic differences in foraging behaviour could be the basis for the genetic influence in the development of feather pecking.

Journal Article↗

Feathers as a means of monitoring mercury in seabirds: Insights from stable isotope analysis.

Mercury concentrations, together with nitrogen and carbon stable isotope signatures, were determined in body feather samples from northern fulmars Fulmarus glacialis and great skuas Catharacta skua, and in different flight feathers from great skuas. There were no significant relationships between trophic status, as defined using isotope analysis, and mercury concentration in the same feather type, in either species. Mercury concentrations in body feather samples were markedly different between fulmars and skuas, reflecting differences in diet, but there was no corresponding difference in trophic status as measured through nitrogen stable isotope signatures. We conclude that mercury concentrations and stable isotope values in feathers are uncoupled, mercury concentrations apparently reflecting the body pool of accumulated mercury at the time of feather growth whilst stable isotope values reflect the diet at the time of feather growth. There were significant positive correlations between the different flight feathers of great skuas for all three parameters measured. These were strongest between primary 10 and secondary 8, suggesting that these two feathers are replaced at the same time in the moult sequence in great skuas. Stable isotope analysis of different feathers may provide a means of investigating moult patterns in birds.

Journal Article↗

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 degrees 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 10 cells per ml) at 50 degrees 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 (10 cells per ml) for 6 days at 50 degrees C. These data indicate a potential biotechnique for degradation and utilization of feather keratin.

Journal Article↗

The effects of Janus Green B on the temporal and spatial pattern of feather germ morphogenesis.

The normal timing and appearance of feather germs was perturbed by injecting the dye Janus Green B into the amniotic fluid of chick embryos at late stage 28, prior to the first appearance of feather germs. This treatment prevented feather germ morphogenesis in some regions while elsewhere it delayed normal morphological development. The Janus Green B effect lasted for approximately 98 hours. Feather regions, which normally form epidermal placodes during the period of treatment, showed the longest delays in subsequent feather germ formation and were the most likely to remain featherless. These results suggest that the epidermal placode stage is critical for feather germ formation. Janus Green B appears to prevent feather germ morphogenesis by interfering with development prior to this critical stage. Since severely affected regions fail to recover their capacity to form feather germs, even after the period of sensitivity to the dye, a limited period of competence is suggested for feather germ formation.

Animals↗

Adaptation to the sky: Defining the feather with integument fossils from mesozoic China and experimental evidence from molecular laboratories.

In this special issue on the Evo-Devo of amniote integuments, Alibardi has discussed the adaptation of the integument to the land. Here we will discuss the adaptation to the sky. We first review a series of fossil discoveries representing intermediate forms of feathers or feather-like appendages from dinosaurs and Mesozoic birds from the Jehol Biota of China. We then discuss the molecular and developmental biological experiments using chicken integuments as the model. Feather forms can be modulated using retrovirus mediated gene mis-expression that mimics those found in nature today and in the evolutionary past. The molecular conversions among different types of integument appendages (feather, scale, tooth) are discussed. From this evidence, we recognize that not all organisms with feathers are birds, and that not all skin appendages with hierarchical branches are feathers. We develop a set of criteria for true avian feathers: 1) possessing actively proliferating cells in the proximal follicle for proximo-distal growth mode; 2) forming hierarchical branches of rachis, barbs, and barbules, with barbs formed by differential cell death and bilaterally or radially symmetric; 3) having a follicle structure, with mesenchyme core during development; 4) when mature, consisting of epithelia without mesenchyme core and with two sides of the vane facing the previous basal and supra-basal layers, respectively; and 5) having stem cells and dermal papilla in the follicle and hence the ability to molt and regenerate. A model of feather evolution from feather bud --> barbs --> barbules --> rachis is presented, which is opposite to the old view of scale plate --> rachis --> barbs --> barbules (Regal, '75; Q Rev Biol 50:35).

Adaptation, Biological↗

The role of mechanical forces on the patterning of the avian feather-bearing skin: A biomechanical analysis of the integumentary musculature in birds.

The integumentary musculature of birds consists of three distinct components. The smooth musculature comprises feather and apterial muscles, which form a continuous musculo-elastic layer within the dermis. The feather muscles, which consistently include at least erectors and depressors, interconnect contour feathers within pterylae (i.e., feather tracts) along gridlines that are oriented diagonally to the longitudinal and transverse axes of the body. The apterial muscles interconnect pterylae by attaching to the contour feathers along their peripheries. The striated musculature is composed of individual subcutaneous muscles, most of which attach to contour feathers along the caudal periphery of pterylae A new integrative functional analysis of the integumentary musculature proposes how apterial muscles stabilize the pterylae and modulate the tension of the musculo-elastic layer, and how subcutaneous muscles provide the initial stimulus for erector muscles being able to ruffle the contour feathers within pterylae. It also shows how the arrangement of the contour feathers and integumentary muscles reflects the stresses and strains that act on the avian skin. These mechanical forces are in effect not only in the adult, especially during flight, but may also be active during feather morphogenesis. The avian integument with its complex structural organization may, therefore, represent an excellent model for analyzing the nature of interactions between the environment and genetic material. The predictions of our model are testable, and our study demonstrates the relevance of integrated analyses of complex organs as mechanically coherent systems for evolutionary and developmental biology.

Animals↗

Bacterial community profiles on feathers during composting as determined by terminal restriction fragment length polymorphism analysis of 16S rDNA genes.

Composting is one of the more economical and environmentally safe methods of recycling feather waste generated by the poultry industry, since 90% of the feather weight consists of crude keratin protein, and feathers contain 15% N. However, the keratin in waste feathers is resistant to biodegradation and may require the addition of bacterial inocula to enhance the degradation process during composting. Two keratin-degrading bacteria isolated from plumage of wild songbirds and identified as Bacillus licheneformis (OWU 1411T) and Streptomyces sp. (OWU 1441) were inoculated into poultry feather composts (1.13 x 10(8) cfu g(-1) feathers) and co-composted with poultry litter and straw in 200-l compost vessels. Composting temperatures, as well as CO(2) and NH(3) evolution, were measured in these vessels to determine the effects of inoculation on the rate and extent of poultry feather decomposition during composting. Terminal restriction fragment length polymorphisms of 16S rRNA genes were used to follow changes in microbial community structure during composting. The results indicated that extensive carbon conversion occurred in both treatments (55.5 and 56.1%). The addition of the bacterial inocula did not enhance the rate of waste feather composting. The microbial community structure over time was very similar in inoculated and uninoculated waste feather composts.

Ammonia↗

Analysis of morphogenesis and keratinization in transfilter recombinants of feather-forming skin.

The relationships between feather morphogenesis, histogenesis, and biochemical differentiation were examined by recombining backskin epidermis and dermis, from chick embryos (Hamburger-Hamilton stages 27-31), with an intervening Nucleopore filter (pore size of 0.4 micron). The filter inhibited normal feather morphogenesis and histogenesis of barb ridges, yet feather-like filaments, which were free of dermal cells, formed from the epidermal cells. Using indirect immunofluorescence, with antiserum against alpha- and beta-keratins, the biochemical differentiation of the feather-like filaments was compared to normal feathers. In the feather-like filaments resulting from tissues of stages 27-29, cells containing beta keratins were occasionally seen at the periphery of the filaments, yet cells containing alpha-keratins were inappropriately located throughout the filaments. In a few feather-like filaments on recombinants resulting from tissues of stages 29.5-31, cells positive for beta-keratins were found in the center of the filament, but again alpha-keratins were also found. Surrounding these cells there were several layers of cells, arranged circumferentially, resembling sheath cells. Some sheath-like cells contained beta-keratins. We conclude that although feather epidermal cells, which are separated from their dermis by a Nuclepore filter, can undergo limited morphogenesis and the production of alpha- and beta-keratins, normal feather morphogenesis, histogenesis, and biochemical differentiation require the intimate associations of epidermis and dermis.

Animals↗

Bird feathers as bioindicators in areas of the German Environmental Specimen Bank--bioaccumulation of mercury in food chains and exogenous deposition of atmospheric pollution with lead and cadmium.

Feathers of selected bird species are used as pollution integrating biomonitors in areas of the German Environmental Specimen Bank. Lead, cadmium and mercury are analysed by Zeeman Solid Sampling AAS in feather vanes of selected bird species with different ecologies. In feather vanes there are quite different distribution patterns of lead and cadmium on the one hand and mercury on the other: lead and cadmium show strong gradients with respect to feather parts with high exposure to atmospheric influences. In contrast to this, mercury is distributed homogeneously. These different distribution patterns of lead and cadmium in feather vanes, in contrast to mercury, show that there are quite different incorporation paths. Cadmium and lead are deposited exogenously and indicate atmospheric pollution. Mercury in feathers is caused endogenously by food and physiology. Most of the mercury analysed in bird's feathers is in the ecotoxic methylated form and the contents in feathers depend on bioaccumulation in food chains. Feathers are suitable indicators for monitoring heavy metal pollution and give us information about incorporation paths and ecotoxic effects.

Aging↗

Apoptosis in feathers of Smyth line chickens with autoimmune vitiligo.

Vitiligo is an acquired dermatological disorder characterized by a loss of epidermal melanocytes resulting in depigmentation of the skin. Mechanisms underlying the destruction of melanocytes in vitiligo remain unclear. An animal model to study spontaneously occurring autoimmune vitiligo is the mutant Smyth line (SL) of chickens. This investigation was designed to determine whether the pathogenesis of depigmentation in Smyth line chicken vitiligo (SLV) involves an apoptotic mechanism. Terminal deoxynucleotide transferase-mediated fluorescein-dUTP nick end labeling (TUNEL) was used to detect in situ cell apoptosis in cryostat sections of 2-week-old regenerating feathers. Two-week-old regenerating feathers were obtained from SL chickens and their normally pigmented controls including the parental Brown line (BL) and Light Brown Leghorn (LBL) chickens at 6, 8, 10 and 12 weeks of age. In feathers from vitiliginous SL chickens, the number of TUNEL+ cells was significantly (P<or=0.05) higher than that in the feathers of non-vitiliginous SL, BL or LBL chickens. These TUNEL+ cells were primarily located in the epithelial barb ridge where melanocyte cell bodies are located. The extent of this apoptosis in the feathers of SLV chickens varied with the severity of depigmentation of the feathers (i.e., highest in active depigmentation), suggesting a close association between apoptosis and the disappearance of melanocytes. In addition to TUNEL staining, most sections were double-stained with monoclonal antibodies specific to either CD8 or MHC class II molecules to further explore the relationship between CD8+ feather-infiltrating lymphocytes and this increase in apoptotic cells. Compared to normally pigmented controls, the number of CD8+ and MHC class II+ cells in the feather pulp and the barb ridge increased 2-4 weeks before the visible onset of SLV, and was directly related to the changes in the number of TUNEL+ cells prior to, at onset and during depigmentation. Moreover, some of these infiltrating CD8+ cells were localized next to or near the TUNEL+ cells. These observations suggest that enhanced apoptosis in the feather of SLV chickens is a pathogenic mechanism involved in the death of melanocytes and appears to be induced by infiltrating cytotoxic T lymphocytes (CD8+).

Animals↗