臺灣博碩士論文加值系統

綠頭鴨 (Anas platyrhynchos) 為肉用之水禽，其雌雄個體之羽毛形態於每年兩次繁殖季節期間（四月至五月；九月至十月），具有兩性異形性（sexual dimorphism）之特徵，但是，雄性綠頭鴨於油壺腺周圍有四支尾羽呈現捲曲狀，與其他尾羽不同，稱之為性捲羽，此四支尾羽則無論是於繁殖季節與否，皆未在雌性綠頭鴨發現此現象。因此，本研究之目的為探討綠頭鴨於成長期間，其性荷爾蒙之變化是否與性捲羽形態之形成相關，並欲探究調控性捲羽形成之分子機制。試驗一，於試驗期間，動物每兩週採血一次取得血清，以ELISA套組 （Cayman Chemical）檢測血清睪固酮 （testosterone, T）與雌二醇 （estradiol, E2）濃度之變化，並配合觀察鴨隻羽毛形態之改變。結果顯示，雌性綠頭鴨於性成熟前E2即開始顯著上升，而雄性綠頭鴨在達性成熟前，T濃度有一個高峰，爾後性捲羽即形成。雄鴨於每年兩次換羽前，即非繁殖羽轉換成繁殖羽（又稱婚羽；pre-nuptial molting）以及繁殖羽轉換成非繁殖羽（post-nuptial molting），其T濃度均會出現一個高峰，且雄鴨轉換成非繁殖羽時，亦伴隨出現一個E2高峰；雌性綠頭鴨於產蛋期前，其E2濃度亦呈現顯著地上升，然而於兩次換羽期間則無明顯濃度改變，其E2濃度水平僅與產蛋期相關。試驗二，將性捲羽與非性捲羽羽毛拔除後，分別於二週（羽毛再生之早期）與四週後（羽毛再生之晚期）採樣取得毛囊，並將毛囊組織分為三個部分，包含真皮乳突（dermal papilla, DP）、外上皮（outside epithelium, O-epi）與毛囊髓質部（pulp, pp）。上述組織於萃取RNA後，利用反轉錄聚合酶連鎖反應（reverse transcription-polymerase chain reaction, RT-PCR）檢測性激素受體，以及羽毛形態生成相關之基因表現。結果顯示，雌性素受體（estrogen receptor α/β, ERα/β）、骨形成蛋白（bone morphogenetic protein 2/4, BMP2/4）與CYP19（aromatase cytochrome P450）於公母鴨之性捲羽與一般尾羽之間的表現並無顯著差異；但雄鴨之性捲羽各部份組織於羽毛再生早期與晚期時，均有雄性素受體（androgen receptor, AR）之表現，但於非性捲羽之尾羽只在DP表現AR。以RNA原位雜交（RNA in situ hybordization）檢測，發現AR表現型式類似RT-PCR結果。試驗三，應用組織免疫化學染色法檢測，發現雌雄鴨之性捲羽與非性捲羽均無表現keratin75（KRT75），而keratin17（KRT17）均表現於羽枝脊（barb ridge）周圍；應用TUNEL套組檢測，發現唯有雄鴨性捲羽之羽軸（rachis）出現細胞凋亡之表現。綜上所述，推測鳥類血清中之性腺激素濃度變化可能與繁殖期、換羽期之啟始，以及羽毛形態有關連性，而雄性綠頭鴨之性捲羽形態可能與雄性素受體（AR）具有一定程度之相關性，但仍需要進一步試驗以瞭解雄性素受體與其他分子之間之調控機制。

Mallard (Anas platyrhynchos) is a common meat-type water fowl. During their breeding seasons (April to May and September to October in Taiwan), the mallard show sexual dimorphic plumage. However, four tail feathers, called sexual curled feathers (SCFs), surrounding the oiler gland in the male mallard display curly morphology, which is different from the rest of tail feathers. Furthermore, the curly-shape feathers exist irrelevantly to the breeding seasons and are absent in the female. Therefore, the aims of this study were to investigate the coordination of changes of serum sex hormone levels and morphogenesis of SCFs during growth, and to study the possible molecular regulation on the morphogenesis of SCFs. In Experiment I, the duck blood samples were collected every two weeks. ELISA kits (Cayman Chemical) were applied to analyze the level of testosterone (T) and estradiol (E2) in mallard serum. Also, the molting phenomenon was recorded during the period of time. Increasing E2 level was noticed in the female before sexual maturity and it reached to the peak after maturity. It was shown that an obviously increased level of T appeared prior to sexual maturity in the male mallard, and at this time point, the SCFs appeared in the male. Interestingly, T peak was found to be associated with molting, i.e. T peak appears before pre-nuptial and post-nuptial molting. Furthermore, it was found that in addition to a T peak, an E2 peak was shown prior to post-nuptial molting. The level of E2 significantly increased before laying period, but no significant difference of E2 level was found between molting in the female mallard, indicating E2 level related to laying only. In Experiment II, two weeks (called early regenerating stage) and four weeks (called late regenerating stage) after plucking SCFs and not curly tail feather (called none sexual curly feathers, nonSCFs) from the male and female mallards, the growing feather follicles were collected and three tissues, including dermal papillae (DP), outside epithelium (O-epi) and pulp (pp), were dissected out for total RNA extraction to analyze the expression of androgen receptor (AR), estrogen receptor α/β (ERα/β), bone morphogenetic protein 2/4 (BMP2/4) and aromatase cytochrome P450 (CYP19) by RT-PCR (reverse transcription-polymerase chain reaction). The result showed that there is no significant difference between SCFs and nonSCFs in the expression of ERα/β, BMP2/4 and CYP19 in all samples. However, AR was found to express in all tissues in SCFs at early and late stages, while it was only expressed in DP in nonSCFs at both stages. The AR expression pattern was consistent with the RNA in situ hybridization results. In Experiment III, no expression of keratin75 (KRT75) was detected in SCFs and nonSCFs in both male and female mallard by immunohistochemistry analysis. However, keratin17 (KRT17) was found to express around barb ridge. The rachis of SCFs from the male mallard showed cell apoptosis by TUNEL assay. In conclusion, the results present in this study have demonstrated that the changes of sex hormone level may be involved in the trigger of breeding, molting and feather morphogenesis. Furthermore, AR, to certain extent, plays an important role on the formation of SCFs. Further studies are required to elucidate the molecular regulation of AR and other molecules on feather morphogenesis.

目錄
前言 1
文獻探討 2
一、 羽毛毛囊構造與羽毛再生週期（Feather follicle and feather regeneration） 2
二、 影響羽毛生長之因子 6
（一） BMP家族與Shh訊息路徑 6
（二） Wnt與β-catenin 訊息路徑 6
（三） FGF家族與Sproutry家族訊息路徑 7
三、 賀爾蒙影響羽毛結構與調節換羽 9
（一） 腎上腺皮質酮（corticosterine） 9
（二） 泌乳素（Prolactin, PRL） 9
（三） 甲狀腺素（Tryriod hormones, THs） 10
四、 性荷爾蒙與第二性徵及羽毛形態之相關性 13
（一） 性荷爾蒙形成路徑 13
（二） 性荷爾蒙影響第二性徵 13
試驗一、建立綠頭鴨之一般血液檢測值 18
一、 前言 18
二、 材料與方法 19
（一） 動物飼養與換羽觀察 19
（二） 血液樣品之採集 19
（三） 動物血清賀爾蒙測定 19
三、 結果 21
（一） 綠頭鴨換羽觀察 21
（二） 綠頭鴨睪固酮（Teststerone, T）濃度趨勢 21
（三） 綠頭鴨血液中雌二醇（Estradiol, E2）濃度趨勢 21
四、 討論 28
試驗二、性賀爾蒙與羽毛形態之相關性 30
一、 前言 30
二、 材料與方法 30
（一） 羽毛毛囊之採集 30
（二） 毛囊RNA之萃取 31
（三） RNA濃度測定 31
（四） 反轉錄聚合酶連鎖反應 （reverse transcription-polymerase chain reaction, RT-PCR） 31
（五） 瓊酯醣凝膠電泳分析與安全染色 33
（六） RNA原位雜合作用(RNA in situ hybordization) 34
三、 結果 37
（一） RT-PCR分析 37
（二） RNA原位雜合分析 37
四、 討論 45
試驗三、影響性捲羽形態結構之因素 47
一、 前言 47
二、 材料與方法 47
（一） 免疫組織化學染色法（immunohistochemistry, IHC） 47
（二） H＆E染色 49
（三） 細胞凋亡檢測（Terminal deoxynucleotidyl transferase dUTP nick end labeling, TUNEL） 50
三、 結果 51
（一） KRT75 51
（二） KRT17 51
（三） TUNEL 51
四、 討論 62
結論 65
參考文獻 66
附錄A-1 80
附錄A-2 81
附錄A-3 82
附錄A-4 83
附錄A-5 84
附錄A-6 85

 
圖次
圖1. 羽毛形態與結構。 4
圖2. 羽毛毛囊結構與羽毛再生週期 5
圖3. 於羽毛再生期間近端處 - 遠端處羽軸與時間維度。 8
圖4. 闡述鳥類和哺乳動物光週期途徑。 12
圖5. 性荷爾蒙合成路徑 15
圖6. E2影響羽毛毛色和ASIP class 1 基因的表現。 16
圖7. 成年岡山土雞鞍部羽毛之兩性異形性與E2對於羽毛結構之影響 17
圖8. 雌雄綠頭鴨之不同部位之羽毛形態 22
圖9. 雄雌綠頭鴨之繁殖羽與褪色羽之羽色差別。 23
圖10. 雄綠頭鴨於不同年齡之羽毛形態。 24
圖11. 綠頭鴨於2015年12月至2016年12月之血清中睪固酮之濃度變化。 25
圖12. 綠頭鴨於2015年12月至2016年12月之血清中雌二醇之濃度變化。 26
圖13. 雄綠頭鴨於2015年12月至2016年12月之血清中雌二醇與睪固酮濃度之變化。 27
圖14. 雄鴨性捲羽以及非性捲羽之毛囊位置示意圖。 38
圖15. 利用RT-PCR分析雄鴨之AR及ERα/β表現。 39
圖16. 利用RT-PCR分析雌鴨之AR及ERα/β表現。 40
圖17. 利用RT-PCR分析雄鴨之CYP19及 BMP2/4表現。 41
圖18. 利用RT-PCR分析雄鴨之CYP19及 BMP2/4表現。 42
圖19. 利用原位雜交分析雄鴨羽毛毛囊內AR之表現。 43
圖20. 利用原位雜交分析雌鴨羽毛毛囊內AR之表現。 44
圖21. 利用免疫組織染色法分析KRT75之表現。 52
圖22. 利用免疫組織染色法分析KRT75之表現。 53
圖23. 利用免疫組織染色法分析KRT17之表現。 54
圖24. 利用免疫組織染色法分析KRT17之表現。 55
圖25. 利用免疫組織染色法分析KRT17之表現。 56
圖26. 利用免疫組織染色法分析KRT17之表現。 57
圖27. 利用免疫組織染色法分析KRT17之表現。 58
圖28. 利用免疫組織染色法分析KRT17之表現。 59
圖29. 利用免疫組織染色法分析TUNEL之表現。 60
圖30. 利用免疫組織染色法分析TUNEL之表現。 61

陳盈豪、曾秋隆、施柏齡、王淑音。2009。去勢對綠頭鴨在婚羽換羽期血球相、血清睪固酮濃度與換羽之影響。台灣獸醫誌35（3）: 217-224。
陳盈豪、王淑音、施柏齡、曾秋隆。2008。去勢對台灣黑土雞在生長期血球相、睪固酮濃度及第二性徵之影響。台灣獸醫誌34（3）: 184-191。
Alibardi, L., and M. Toni. 2008. Cytochemical and molecular characteristics of the process of cornification during feather morphogenesis. Prog Histochem Cytochem 43:1-69.
Andrews, J. E., C. A. Smith, and A. H. Sinclair. 1997. Sites of estrogen receptor and aromatase expression in the chicken embryo. Gen. Comp. Endocrinol. 108:182-190.
Beek, .V., N., E. Bodo, A. Kromminga, E. Gaspar, K. Meyer, M. A. Zmijewski, A. Slominski, B. E. Wenzel, and R. Paus. 2008. Thyroid hormones directly alter human hair follicle functions: anagen prolongation and stimulation of both hair matrix keratinocyte proliferation and hair pigmentation. J. Clin. Endocrinol. Metab. 93:4381-4388.
Bentlley, G.E. 2008. Biological timing: sheep, Dr. Seuss, and mechanistic ancestry. Curr Biol. 18:736-738.
Billoni, N., B. Buan, B. Gautier, O. Gaillard, Y. F. Mahe, and B. A. Bernard. 2000. Thyroid hormone receptor beta1 is expressed in the human hair follicle. Br. J. Dermatol. 142:645-652.
Brake, J., P. Thaxton, and E. H. Benton. 1979. Physiological changes in caged layers during a forced molt. 3. Plasma thyroxine, plasma triiodothyronine, adrenal cholesterol, and total adrenal steroids. Poultry science 58:1345-1350.
Brian, W. M., D. Wang, A. M. Noone, N. Liu, N. Harazduk, M. Lumpkin, A. Haramati, P. Saunders, M. A. Dutton, and H. Amri. 2011. Stress biomarkers in medical students participating in a mind body medicine skills program. Evid. Based Complement. Alternat. 2011:950461.
Bruggeman, V., P. Van As, and E. Decuypere. 2002. Developmental endocrinology of the reproductive axis in the chicken embryo. Comp. Biochem. Physiol. A. Mol. Integr. Physiol. 131:839-846.
Burke, W. H., and M. H. Henry. 1999. Gonadal development and growth of chickens and turkeys hatched from eggs injected with an aromatase inhibitor. Poult. Sci. 78:1019-1033.
Carson, C., 3rd, and R. Rittmaster. 2003. The role of dihydrotestosterone in benign prostatic hyperplasia. Urology 61:2-7.
Chen, C. C., M. V. Plikus, P. C. Tang, R. B. Widelitz, and C. M. Chuong. 2015. The modulatable stem cell niche: Tissue interactions during hair and feather follicle regeneration. J. Mol. Biol.
Cho, Y.M., Woodard, G.L., Dunbar, M., Gocken, T., Jimenez, J.A., and Foley, J. 2003. Hair-cycle-dependent expression of parathyroid hormone-related protein and its type I receptor: Evidence for regulation at the anagen to catagen transition. J. Invest. Dermatol. 120: 715-727.
Chodankar, R., C. H. Chang, Z. Yue, T. X. Jiang, S. Suksaweang, L. Burrus, C. M. Chuong, and R. Widelitz. 2003. Shift of localized growth zones contributes to skin appendage morphogenesis: role of the Wnt/beta-catenin pathway. J. Invest. Dermatol. 120:20-26.
Contreras-Jurado, C., L. Garcia-Serrano, M. Martinez-Fernandez, L. Ruiz-Llorente, J. M. Paramio, and A. Aranda. 2014. Impaired hair growth and wound healing in mice lacking thyroid hormone receptors. PloS one 9:e108137.
Dawson, A. 2005. The effect of temperature on photoperiodically regulated gonadal maturation, regression and moult in starlings-potential consequences of climate change. Funct. Ecol. 19:995-1000.
Dawson, A. 2006. Control of molt in birds: association with prolactin and gonadal regression in starlings. Gen. Comp. Endocrinol. 147:314-322.
Dawson, A., and A. R. Goldsmith. 1982. Prolactin and gonadotrophin secretion in wild starlings （Sturnus vulgaris） during the annual cycle and in relation to nesting, incubation, and rearing young. Gen. Comp. Endocrinol. 48:213-221.
Dawson, A., C. M. Perrins, P. J. Sharp, D. Wheeler, and S. Groves. 2009. The involvement of prolactin in avian molt: the effects of gender and breeding success on the timing of molt in Mute swans （Cygnus olor）. Gen. Comp. Endocrinol. 161:267-270.
Dawson, A., V. M. King, G. E. Bentley, and G. F. Ball. 2001. Photoperiodic control of seasonality in birds. J. Biol. Rhythms. 16:365-380.
Day, L. B., J. T. McBroom, and B. A. Schlinger. 2006. Testosterone increases display behaviors but does not stimulate growth of adult plumage in male golden-collared manakins (Manacus vitellinus). Horm. Behav. 49:223-232.
DesRochers, D. W., J. M. Reed, J. Awerman, J. A. Kluge, J. Wilkinson, L. I. Griethuijsen, J. Aman, and L. M. Romero. 2009. Exogenous and endogenous corticosterone alter feather quality. Comp. Biochem. Physiol. A 152:46-52.
Donham, R. S. 1979. Annual cycle of plasma luteinizing hormone and sex hormones in male and female mallards （Anas platyrhynchos）. Biol. Reprod. 21:1273-1285.
Dunn, P. O., L. A. Whittingham, and T. E. Pitcher. 2001. Mating systems, sperm competition, and the evolution of sexual dimorphism in birds. Evolution 55:161-175.
Eikenaar, C., M. Whitham, J. Komdeur, M. V. D. Velde, and I. T. Moore. 2011. Testosterone, plumage colouration and extra-pair paternity in male north- American barn swallows. PLoS ONE 6:e23288.
Foitzik, K., G. Lindner, S. Mueller-Roever, M. Maurer, N. Botchkareva, V. Botchkarev, B. Handjiski, M. Metz, T. Hibino, T. Soma, G. P. Dotto, and R. Paus. 2000. Control of murine hair follicle regression （catagen） by TGF-beta1 in vivo. FASEB J. 14:752-760.
Frankenhuis, M. T., and H. J. Kappert. 1980. Experimental transformation of right gonads of female fowl in to fertile testes. Biol. Reprod. 23:526-529.
Freinkel, R. K., and N. Freinkel. 1972. Hair growth and alopecia in hypothyroidism. Arch. Dermatol. 106:349-352.
Gelmann, E. P. 2002. Molecular biology of the androgen receptor. J. Clin. Oncol. 20:3001-3015.
George, F. W., and J. D. Wilson. 1980. pathogenesis of the henny feathering trait in the Sebright bantam chicken. Increased conversion of androgen to estrogen in skin. J. Clin. Invest. 66:57-65.
Gonzalez, G., G. Sorci, L. C. Smith, and F. D. Lope. 2001. Testosterone and sexual signalling in male house sparrows (Passer domesticus). Behav. Ecol. Sociobiol. 50:557-562.
Greenwold, M.J., R. H. Sawyer. 2013. Molecular evolution and expression of archosaurian beta-keratins: diversification and expansion of archosaurian beta-keratins and the origin of feather beta-keratins. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 320, 393-405.
Haase, E. 1983. The annual reproductive cycle in mallards. J. Steroid Biochem. 19:731-737.
Haase, E., and R. Schmedemann. 1992. Dose-dependent effect of testosterone on the introduction of eclipse coloration in castrated wild mallard drake （Anas platyrhynchos.） Can. J. Zool. 70:428-431.
Haase, E., P. J. Sharp, and E. Paulke. 1985. Seasonal changes in the concentrations of plasma gonadotropins and prolactin in wild mallard drakes. J. Exp. Zool. 234:301-305.
Haase, E., S. Ito, and K. Wakamatsu. 1995. Influences of sex, castration, and androgens on the eumelanin and pheomelanin contents of different feathers in wild mallards. Pigm. Cell Res. 8:164-170.
Hacohen, N., S. Kramer, D. Sutherland, Y. Hiromi, M. A. Krasnow. 1998. sprouty encodes a novel antagonist of FGF signaling that patterns apical branching of the Drosophila airways. Cell 92, 253–263.
Hagelin, J. C. 2001. Castration in Gambel's and Scaled Quail: ornate plumage and dominance persist, but courtship and threat behaviors do not. Horm. Behav. 39:1-10.
Hagelin, J. C., and R. T. Kimball. 1997. A female Gambel's quail in partial male plumage. Wilson Bull. 109: 544-546.
Hasse, E., R. Schmedemann. 1992. Dose-dependent effect of testosterone on the introduction of eclipse coloration in castrated wild mallard duck (Anas platyrhynchos). Can J Zoll 70:428-431.
Hebert, J. M., T. Rosenquist, J. Gotz, and G. R. Martin. 1994. FGF5 as a regulator of the hair growth cycle: Evidence from targeted and spontaneous mutations. Cell 78: 1017-1025.
Herremans, M., L. Berghman, E. Decuypere, and F. Vandesande. 1993. Immunocytochemical demonstration of progesterone and estrogen receptors in feathers and skin of adult hens. Acta. Biol. Hung. 44:353-366.
Hesse, M., A. Zimek, K. Weber, T. M. Magin. 2004. Comprehensive analysis of keratin gene clusters in humans and rodents. Eur J Cell Biol 83: 19-26.
Hibberts, N. A., A. E. Howell, and V. A. Randall. 1998. Balding hair follicle dermal papilla cells contain higher levels of androgen receptors than those from non-balding scalp. J. Endocrinol. 156:59-65.
Hickman, C. P., L. S. Roberts, A. L. Larson. (2003). Integrated principles of zoology. Dubuque, IA: McGraw-Hill. P.538. ISBN 0-07-243940-8.
Hida, T., K. Wakamatsu, E. V. Sviderskaya, A. J. Donkin, L. Montoliu, L. M. Lynn, B. Yu, G. L. Millhauser, S. Ito, G. S. Barsh, K. Jimbow, and D. C. Bennett. 2009. Agouti protein, mahogunin, and attractin in pheomelanogenesis and melanoblast-like alteration of melanocytes: a cAMP-independent pathway. Pigment Cell Melanoma Res. 22(5):623-634.
Hirota, R., S. Tajima., Y. Yoneda, T. Tamayama, M. Watanabe, K. Ueda, T. Kubota, and R. Yoshida. 2002. Alopecia of IFN-γ knockout mouse as a model for disturbance of the hair cycle: A unique arrest of the hair cycle at the anagen phase accompanied by mitosis. J. Interferon Cytokine Res. 22: 935–945.
Huelsken, J., R. Vogel, B. Erdmann, G. Cotsarelis, and W. Birchmeier. 2001. beta-Catenin controls hair follicle morphogenesis and stem cell differentiation in the skin. Cell 105:533-545.
Ikegami, K., and T. Yoshimura. 2012. Circadian clocks and measurement of daylength in seasonal reproduction. Mol Cell Endocrinol 349 (1):76-81.
Inui, S., Y. Fukuzato, T. Nakajima, K. Yoshikawa, and S. Itami. 2002. Androgen-inducible TGF-β1 from balding dermal papilla cells inhibits epithelial cell growth: a clue to understanding paradoxical effects of androgen on human hair growth. FASEB J. 16:1967-1969.
Jamora, C., DasGupta, R., Kocieniewski, P., and Fuchs, E. 2003. Links between signal transduction, transcription and adhesion in epithelial bud development. Nature 422: 317–322.
Kimball RT., J. D. Ligon. 1999. Evolution of avain plumge dichromatism from a proximate perspective. Am Nat 154:182-193.
Kishimoto, J., R. E. Burgeson, and B. A. Morgan. 2000. Wnt signaling maintains the hair-inducing activity of the dermal papilla. Genes. Dev. 14:1181-1185.
Kowata, K., M. Nakaoka, K. Nishio, A. Fukao, A. Satoh, M. Ogoshi, S. Takahashi, M. Tsudzuki, and S. Takeuchi. 2014. Identification of a feather β-keratin gene exclusively expressed in pennaceous barbule cells of contour feathers in chicken. Gene. 542(1):23-28.
Kuenzel, W. J. 2003. Neurobiology of molt in avian species. Poult. Sci. 82:981-991.
Kuenzel, W. J., and C. W. Helms. 1974. An annual cycle study of tan-striped and white-striped White-throated sparrows. Auk 91:44-53.
Lahaye, S. E., M. Eens, V. M. Darras, and R. Pinxten. 2014. Bare-part color in female budgerigars changes from brown to structural blue following testosterone treatment but is not strongly masculinized. PLoS One. 9(1):e86849.
Leiros, G. J., A. I. Attorresi, and M. E. Balana. 2012. Hair follicle stem cell differentiation is inhibited through cross-talk between Wnt/beta-catenin and androgen signalling in dermal papilla cells from patients with androgenetic alopecia. Br. J. Dermatol. 166:1035-1042.
Leska, A., J. Kiezun, B. Kaminska, and L. Dusza. 2015. Estradiol concentration and the expression of estrogen receptors in the testes of the domestic goose （Anser anser f. domestica） during the annual reproductive cycle. Domest. Anim. Endocrinol. 51:96-104.
Li, J. J., L. H. Mitchell, and R. L. Dow. 2010. Thyroid receptor agonists for the treatment of androgenetic alopecia. Bioorg. Med. Chem. Lett. 20:306-308.
Ligon, J. D., and P. W. Zwartjes. 1995. Ornate plumage of male red junglefowl does not influence mate choice by females. Anim. Behav. 49: 117-125.
Ligon, J. D., R. Thornhill, M. Zuk, and K. Johnson. 1990. Male-male competition, ornamentation and the role of testosterone in sexual selection in red jungle fowl. Anim. Behav. 40: 367-373.
Lin, C. M., T. X. Jiang, R. B. Widelitz, and C. M. Chuong. 2006. Molecular signaling in feather morphogenesis. Curr. Opin. Cell. Biol. 18:730-741.
Lin, S. J., R. B. Widelitz, Z. Yue, A. Li, X. Wu, T. X. Jiang, P. Wu, and C. M. Chuong. 2013. Feather regeneration as a model for organogenesis. Dev. Growth Differ. 55:139–148.
Lincoln, G. A. 1971. Puberty in a seasonally breeding male, the red deer stag （Cervus elaphus L.）. J. Reprod. Fertil. 25:41-54.
Lindsay, W. R., M. S. Webster, and H. Schwabl. 2011. Sexually selected male plumage color is testosterone dependent in a tropical passerine bird, the red-backed fairy-wren (Malurus melanocephalus). PLoS ONE 6: e26067. doi:10.1371/journal.pone.0026067.
Lu, D., D. Willard, I. R. Patel, S. Kadwell, L. Overton, T. Kost, M. Luther, W. Chen, R. P. Woychik, and W. O. Wilkison. 1994. Agouti protein is an antagonist of the melanocyte-stimulating-hormone receptor. Nature. 371(6500):799-802.
Maia, R., J. V. O. Caetano, S. N. Bao, and R. H. Macedo. 2009. Iridescent structural colour production in male blue-black grassquit feather barbules: the role of keratin and melanin. J. R. Soc. Interface. 6:S203-S211.
Massague, J., and Y. G. Chen. 2000. Controlling TGF-beta signaling. Genes Dev. 14:627-644.
Mayer, J. A., C. M. Chuong, and R. Widelitz. 2004. Rooster feathering, androgenic alopecia, and hormone-dependent tumor growth: what is in common? Differentiation; research in biological diversity 72:474-488.
McElwee, K.J., S. Kissling, E. Wenzel, A. Huth, R. Hoffmann. 2003 Cultured peribulbar dermal sheath cells can induce hair follicle development and contribute to the dermal sheath and dermal papilla. J Invest Dermatol. 121:1267–1275.
McGowan, K. M., P. A. Coulombe. 1998b. The wound repair associated keratins 6, 16, and 17: Insights into the role of intermediate filaments in specifying cytoarchitecture. In: Harris JR, Herrmann H, editors. Subcellular biochemistry: Intermediate filaments. London, UK: Plenum Publishing. pp. 141-165.
McGowan, K., P. A. Coulombe. 1998a. Onset of keratin 17 expression coincides with the definition of major epithelial lineages during mouse skin development. J Cell Biol. 143:469-486.
Millar, S. E. 2002. Molecular mechanisms regulating hair follicle development. J. Invest. Dermatol. 118:216-225.
Mundy, N. I. 2005. A window on the genetics of evolution: MC1R and plumage colouration in birds. Proc Biol Sci. 272(1573):1633-40.
Mundy, N. I., N. S. Badcock, T. Hart, K. Scribner, K. Janssen, andN. J. Nadeau. 2004. Conserved genetic basis of a quantitative plumage trait involved in mate choice. Science. 303(5665):1870-3.
Ng, C. S., P. Wu, J. Foley, A. Foley, M. L. McDonald, W. T. Juan, C. J. Huang, Y. T. Lai, W. S. Lo, C. F. Chen, S. M. Leal, H. Zhang, R. B. Widelitz, P. I. Patel, W. H. Li, and C. M. Chuong. 2012. The chicken frizzle feather is due to an α-keratin (KRT75) mutation that causes a defective rachis. PLoS Genet. 8:e1002748.
Ohnemus, U., M. Uenalan, J. Inzunza, J. A. Gustafsson, and R. Paus. 2006. The hair follicle as an estrogen target and source. Endocr. Rev. 27:677-706.
Oribe, E., A. Fukao, C. Yoshihara, M. Mendori, K. G. Rosal, S. Takahashi, and S. Takeuchi. 2012. Conserved distal promoter of the agouti signaling protein （ASIP） gene controls sexual dichromatism in chickens. Gen. Comp. Endocrinol. 177:231-237.
Ouji, Y., M. Yoshikawa, K. Moriya, M. Nishiofuku, R. Matsuda, and S. Ishizaka. 2008. Wnt-10b, uniquely among Wnts, promotes epithelial differentiation and shaft growth. Biochem. Biophys. Res. Commun. 367:299-304.
Paulke, E., and E. Haase. 1978. A comparison of seasonal changes in the concentrations of androgens in the peripheral blood of wild and domestic ducks. Gen. Comp. Endocrinol. 34:381-390.
Paus, R. and Foitzik, K. 2004. In search of the ‘hair cycle clock’: A guided tour. Differentiation 72: 489–511.
Plikus, M. V., J. A. Mayer, D. de la Cruz, R. E. Baker, P. K. Maini, R. Maxson, and C. M. Chuong. 2008. Cyclic dermal BMP signalling regulates stem cell activation during hair regeneration. Nature 451:340-344.
Poppe, K., B. Velkeniers, and D. Glinoer. 2007. Thyroid disease and female reproduction. Clin. Endocrinol. （Oxf） 66:309-321.
Queen, W. H., V. L. Christensen, and J. D. May. 1997. Supplemental thyroid hormones and molting in turkey breeder hens. Poultry science 76:887-893.
Roberts, M. L., E. Ras, and A. Peters. 2009. Testosterone increases UV reflectance of sexually selected crown plumage in male blue Tits. Behav. Ecol. 20:535-541.
Rogers, G. E. 1984. Studies on keratinmultigene families. Biochemical Society Symposium. 49, 85-108.
Romero, L. M., and L. R. Healey. 2000. Daily and seasonal variation in response to stress in captive starlings (Sturnus vulgaris): corticosterone. Gen. Comp. Endocrinol. 119: 52-59.
Romero, L. M., D. Strochlic, and J. C. Wingfield. 2005. Corticosterone inhibits feather growth: potential mechanism explaining seasonal down regulation of corticosterone during molt. Comp. Biochem. Physiol. A 142:65-73.
Romero, L. M., N. E. Cry, and R. C. Romero. 2006. Corticosterone responses change seasonally in free-living house sparrows (Passer domesticus). Gen. Comp. Endocrinol. 149, 58-65.
Safer, J. D., L. M. Fraser, S. Ray, and M. F. Holick. 2001. Topical triiodothyronine stimulates epidermal proliferation, dermal thickening, and hair growth in mice and rats. Thyroid 11:717-724.
Saino, N., and A. P. Moller. 1994. Secondary sexual characters, parasites, and testosterone in the barn swallow, Hirundo rustica. Anim. Behav. 48:1325-1333.
Scheib, D. 1983. Effects and role of estrogens in avian gonadal differentiation. Differentiation; research in biological diversity 23 Suppl:S87-92.
Schleussner, G., J. P. Dittami, and E. Gwinner. 1985. Testosterone implants affect molt in male European starlings, Sturnus vulgaris. Physiol. Zool. 58:597-604.
Sechman, A., K. Pawlowska, and J. Rzasa. 2009. Influence of triiodothyronine （T（3）） on secretion of steroids and thyroid hormone receptor expression in chicken ovarian follicles. Domest. Anim. Endocrinol. 37:61-73.
Sharp, P. J. 2005. Photoperiodic regulation of seasonal breeding in birds. Ann. N. Y. Acad. Sci. 1040:189-199.
Sharp, P. J., A. Dawson, and R. W. Lea. 1998. Control of luteinizing hormone and prolactin secretion in birds. Comp. Biochem. Physiol. C. Pharmacol. Toxicol. Endocrinol. 119:275-282.
Shimizu, H., and B. A. Morgan. 2004. Wnt signaling through the beta-catenin pathway is sufficient to maintain, but not restore, anagen-phase characteristics of dermal papilla cells. J. Invest. Dermatol. 122:239-245.
Siefferman, L., M. Liu, K. J. Navara, M. T. Mendonça, and G. E. Hill. 2013. Effect of prenatal and natal administration of testosterone on production of structurally based plumage coloration. Physiol. Biochem. Zool. 86:323-332.
Slebodzinski, A. B. 2005. Ovarian iodide uptake and triiodothyronine generation in follicular fluid. The enigma of the thyroid ovary interaction. Domest. Anim. Endocrinol. 29:97-103.
Sockman, K. W., P. J. Sharp, and H. Schwabl. 2006. Orchestration of avian reproductive effort: an integration of the ultimate and proximate bases for flexibility in clutch size, incubation behaviour, and yolk androgen deposition. Biol. Rev. Camb. Philos. Soc. 81:629-666.
Soma, T., Y. Tsuji, and T. Hibino. 2002. Involvement of transforming growth factor-beta2 in catagen induction during the human hair cycle. J. Invest. Dermatol. 118:993-997.
Somes, R. G. 1990. Mutations and major variants of plumage and skin in chickens. In: Crawford RD, editor. Poultry Breeding and Genetics. Amsterdam, Netherlands: Elsevier. pp. 169–208.
Strochlic, D. E., and L. M. Romero. 2008. The effects of chronic psychological and physical stress on feather replacement in European starlings (Sturnus vulgaris). Comp. Biochem. Physiol. A 149:68-79.
Suzuki, H. A., T. Sato, H. Nakamura. 2005. Regulation of isthmic Fgf8 signal by sprouty2. Development 132, 257–265.
Thapliyal, A., A. Chandola-Saklani, D. Bhatt, and P. Anthwal. 2014. Binding pattern of 125iodine thyroxine and tri-iodothyronine in skin and liver tissues of spotted munia, Lonchura punctulata: co-relation to seasonal cycles of breeding and molting. Indian J Exp Biol 52:478-488.
Thomas, F. S., and S. A. Kay. 2003. Circadian clocks in daily and seasonal control of development. Science 301: 326-330.
Tonra, C. M., K. L. D. Marini, P. P. Marra, R. R. Germain, R. L. Holberton, and M. W. Reudink. 2014. Color expression in experimentally regrown feathers of an overwintering migratory bird: implications for signaling and seasonal interactions. Ecol. Evol. 4:1222-1232.
Troyanovsky, S. M., V. I. Guelstein, T. A. Tchipysheva, V. A rutovskikh, G. A. annikov. Patterns of expression of keratin 17 in human epithelia: Dependency on cell position. J Cell Sci. 1989;93:419–426.
Truica, C. I., S. Byers, and E. P. Gelmann. 2000. Beta-catenin affects androgen receptor transcriptional activity and ligand specificity. Cancer Res. 60:4709-4713.
Widelitz, R. B., T. X. Jiang, C. W. Chen, N. S. Stott, H. S. Jung, and C. M. Chuong. 1999. Wnt-7a in feather morphogenesis: involvement of anterior-posterior asymmetry and proximal-distal elongation demonstrated with an in vitro reconstitution model. Development 126:2577-2587.
Widelitz, R. B., T. X. Jiang, J. Lu, and C. M. Chuong. 2000. beta-catenin in epithelial morphogenesis: conversion of part of avian foot scales into feather buds with a mutated beta-catenin. Dev. Biol. 219:98-114.
Woods, J. E., and L. V. Domm. 1966. A histochemical identification of the androgen-producing cells in the gonads of the domestic fowl and albino rat. Gen. Comp. Endocrinol. 7:559-570.
Yang, X., J. Zheng, R. Na, J. Li, G. Xu, L. Qu, and N. Yang. 2008. Degree of sex differentiation of genetic female chicken treated with different doses of an aromatase inhibitor. Sex. Dev. 2:309-315.
Yoshihara, C., A. Fukao, K. Ando, Y. Tashiro, S. Taniuchi, S. Takahashi, and S. Takeuchi. 2012. Elaborate color patterns of individual chicken feathers may be formed by the agouti signaling protein. Gen Comp Endocrinol. 175(3):495-9.
Yoshimura, T., S. Yasuo, M. Watanabe, M. Iigo, T. Yamamura, K. Hirunagi, and S. Ebihara. 2003. Light-induced hormone conversion of T4 to T3 regulates photoperiodic response of gonads in bird. Nature 426:178-181.
Yu, M. K., Z. C. Yue, P. Wu, D. Y. Wu, J. A. Mayer, M. Medina, R. B. Widelitz, T. X. Jiang, and C. M. Chuong. 2004b. The developmental biology of feather follicles. Int. J. Dev. Biol. 48:181-191.
Yu, M., P. Wu, R. B. Widelitz, and C. M. Chuong. 2002. The morphogenesis of feathers. Nature 420:308-312.
Yu, M., Z. Yue, P. Wu, D. Y. Wu, J. A. Mayer, M. Medina, R. B. Widelitz, T. X. Jiang, and C. M. Chuong. 2004a. The biology of feather follicles. Int. J. Dev. Biol. 48:181-191.
Yue, Z., T. X. Jiang, P. Wu, R. B. Widelitz, and C. M. Chuong. 2012. Sprouty/FGF signaling regulates the proximal–distal feather morphology and the size of dermal papillae. Dev. Biol. 372:45-54.
Yue, Z., T. X. Jiang, R. B. Widelitz, and C. M. Chuong. 2005. Mapping stem cell activities in the feather follicle. Nature 438:1026-1029.
Yue, Z., T. X. Jiang, R. B. Widelitz, and C. M. Chuong. 2006. Wnt3a gradient converts radial to bilateral feather symmetry via topological arrangement of epithelia. Proc. Natl. Acad. Sci.103:951-955.
Yue, Z., T. X. Jiang, R. B. Widelitz, and C. M. Chuong. 2006. Wnt3a gradient converts radial to bilateral feather symmetry via topological arrangement of epithelia. PNAS 103:951-955.