簡易檢索 / 詳目顯示

研究生: 洪心怡
Hung, Hsin-Yi
論文名稱: 類胡蘿蔔素呈色及黑色素呈色的特徵在紅嘴黑鵯的研究
Studies of carotenoid-based and melanin-based characteristics in Himalayan black bulbul (Hypsipetes leucocephalus regerrimus)
指導教授: 李壽先
Li, Shou-Hsien
學位類別: 博士
Doctor
系所名稱: 生命科學系
Department of Life Science
論文出版年: 2015
畢業學年度: 103
語文別: 英文
論文頁數: 118
中文關鍵詞: 紅嘴黑鵯類胡蘿蔔素黑色素性擇鳥類通訊
英文關鍵詞: Himalayan black bulbul, Hypsipetes leucocephalus regerrimus, carotenoid-based trait, melanin-based trait, quality cue
論文種類: 學術論文
相關次數: 點閱:73下載:14
分享至:
查詢本校圖書館目錄 查詢臺灣博碩士論文知識加值系統 勘誤回報
  • 鳥類體表顏色多樣性極大,這些顏色大部分具有功能性,對於鳥類溝通非常重要。在各種體表顏色中,由類胡蘿蔔素呈色 (呈現黃、橘或紅色)以及黑色素呈色(黑、灰或棕色)的特徵是最普遍也最重要的特徵。這類特徵通常能傳遞個體的訊息,並且具有重要的功能,例如個體可據此選擇配偶或評估對手。傳統上,類胡蘿蔔素呈色的特徵,被普遍認為是個體選擇配偶的依據,而黑色素呈色的特徵,則被認為與社會互動有關。但是這種觀念已被質疑,因為隨著越來越多的鳥種被研究,發現這兩種特徵在不同種類,可能功能不同,顯示這兩種特徵在鳥類的演化機制,可能比目前所知的更為複雜。除此之外,大部分的相關研究都在探討這兩類特徵在性擇上扮演的角色,所以多是以兩性有明顯型態差異的物種為研究標的(這類物種通常可能受到較高的性擇壓力),較少以在兩性型態差異較小的物種為主題,而這些物種可能受到與兩性型態差異大的物種不同的選汰壓力。為了能更完整地了解顏色在鳥類溝通上的演化機制,需要研究這兩類特徵在兩性型態差異較小的物種上扮演的角色。本研究選擇紅嘴黑鵯 (Hypsipetes leucocephalus nigerrimus )為研究物種,他們具有類胡蘿蔔素呈色的喙部、跗蹠與黑色素呈色的羽毛,對人眼來說是雌雄單型性的鳥種。我的結果顯示,在鳥類視覺上,這兩類特徵在兩性間是有差異的。而類胡蘿蔔素呈色的喙部與黑色素呈色的胸部與肩部羽毛的顏色表現,也與個體的生理狀況(包括個體的免疫能力與受到的氧化壓力)有關。雌性偏好試驗的結果顯示,單獨類胡蘿蔔素呈色的部位,可能不是雌性黑鵯選擇配偶的依據。我的論文提出清楚的證據,顯示類胡蘿蔔素呈色以及黑色素呈色的特徵,都能顯示紅嘴黑鵯個體的生理狀況,因此對紅嘴黑鵯而言,這些特徵可能具有重要的功能。但是要知道這些特徵在這個物種真正的功能,還需要之後更詳細的研究。

    Integumentary colorations are essential signals in avian communication. Among all the various color types, carotenoid-based (expressed in yellow, orange or red) and melanin-based (expressed in black, grey or brown) colors are the most common and important traits in birds. Studies have shown that these traits are informative and serve vital functions, such as cues for choosing mates or assessing opponents. Traditionally, in avian communication, carotenoid-based colors are thought to be sexually selected traits and melanin-based colors signal social status. However, the results of various studies indicate these traits serve several functions in different avian species. It suggests that the evolution of both traits is more complex than we used to think. Nevertheless, most relevant studies have focused on the role in sexual dichromatic species, which might be suggested to be under higher pressure of sexual selection. There is a gap of how these traits evolved and what their functions are in less sexually dichromatic species, which might be under different selection regimes from sexually dichromatic ones. In order to understand the imperatives of coloration in avian communication completely, studies to characterize variations and functionality of both traits on less sexually dichromatic species are needed. I chose to study Himalayan black bulbuls (Hypsipetes leucocephalus nigerrimus), which are sexually monomorphic to human vision and contain a carotenoid-based bill and tarsi and melanin-based plumage to analyze the possible roles of both types of traits. My results showed that both carotenoid- and several melanin-based parts were sexually dichromatic in avian vision. Furthermore, I found that the expression of a carotenoid-based bill and melanin-based breast and scapular colorations were correlated with individuals’ physical conditions, including immunocompetence and oxidative stress levels. The results of the female-preference test on carotenoid-based traits showed that the red bared parts alone may not be the cues for mate choice for female black bulbuls. My dissertation provides clear evidence that both carotenoid- and melanin-based traits should be informative cues reflecting bearers’ physical condition; it suggests that these traits may play a role in the signaling of Himalayan black bulbuls, but the functions of the traits need to be further investigated.

    Table of contents 1 List of Tables 2 List of Figures 3 致謝 4 Abstract 7 中文摘要 9 Chapter One General introduction 11 Chapter Two Sexual dichromatism cryptic to the human eye and the quality of museum specimens in the Himalayan black bulbul (H. leucocephalus nigerrimus, Pycnonotidae) 23 Chapter Three Negative effects of molting on carotenoid-based characteristics and individual’s physical condition in the Himalayan black bulbul (H. leucocephalus nigerrimus) 52 Chapter Four Negative correlation between melanin-based plumage coloration and oxidative stress in Himalayan black bulbul (H. leucocephalus nigerrimu) 73 Chapter Five Carotenoid-based traits alone might not be the sexually selected cues for female Himalayan black bulbuls (H. leucocephalus nigerrimu) 90 Chapter Six General conclusions and perspectives 105 Supplementary materials 115

    Chapter One
    Alonso‐Alvarez C, Bertrand S, Devevey G, Gaillard M, Prost J, Faivre B, Sorci G (2004) An experimental test of the dose‐dependent effect of carotenoids and immune activation on sexual signals and antioxidant activity. Am Nat 164:651-659.
    Andersson M, Simmons L W (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296-302.
    Badyaev A V, Hill G E (2000) Evolution of sexual dichromatism: contribution of carotenoid‐versus melanin‐based coloration. Bio J Linn Soci 69:153-172.
    Bize P, Gasparini J, Klopfenstein A, Altwegg R, Roulin A (2006) Melanin based coloration is a nondirectionally selected sex specific signal of offspring development in the alpine swift. Evolution 60:2370-2380.
    Blount J D, Matheson S M (2006) Effects of carotenoid supply on escape flight responses in zebra finches, Taeniopygia guttata. Anim Behav 72:595-601.
    Bortolotti G R (2006) Natural selection and coloration: protection, concealment, advertisement, or deception. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard Universtiy Press, United Kingdom.
    Bostrom M R, Ritchison G (2006) Possible relationships between morphology, territory quality, and skin color of American Kestrels1. J Field Ornithol 77:392-398.
    Brush A H (1990) Metabolism of carotenoid pigments in birds. FASEB J 4:2969.
    Casagrande S, Csermely D, Pini E, Bertacche V, Tagliavini J (2006) Skin carotenoid concentration correlates with male hunting skill and territory quality in the kestrel Falco tinnunculus. J Avian Biol 37:190-196.
    Chew B P (1996) Importance of antioxidant vitamins in immunity and health in animals. Anim Feed Sci Technol 59:103-114.
    Chew B P, Park J S (2004) Carotenoid action on the immune response. J Nutr 134:257S-261S.
    Da Silva A, Van Den Brink V, Emaresi G, Luzio E, Bize P, Dreiss A N, Roulin A (2013) Melanin-based colour polymorphism signals aggressive personality in nest and territory defence in the tawny owl (Strix aluco). Behav Ecol Sociob 67:1041-1052.
    Dale J (2000) Ornamental plumage does not signal male quality in red-billed queleas. Proc R Soc Lond B Biol Sci 267:2143-2149.
    Dale J (2006) Intraspecific Variation in Coloration. In: Hill and McGraw (ed) Bird coloration: function and evolution 3rd edn. Harvard University Press, United Kingdom.
    Faivre B, Gregoire A, Preault M, Cezilly F, Sorci G (2003) Immune activation rapidly mirrored in a secondary sexual trait. Science 300:103.
    Fargallo J A, Laaksonen T, Korpimaki E, Wakamatsu K (2007) A melanin-based trait reflects environmental growth conditions of nestling male Eurasian kestrels. Evol Ecol 21:157-171.
    Fox D L (1976) Animal biochromes and structural colours: physical, chemical, distributional & physiological features of coloured bodies in the animal world. Univ of California Press, USA.
    Gray D A (1996) Carotenoids and sexual dichromatism in North American passerine birds. Am Nat 453-480.
    Hill G E (1991) Plumage coloration is a sexually selected indicator of male quality. Nature 350:337-339.
    Hill G E (2006) Female mate choice for ornamental coloration. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard Universtiy Press, United Kingkom.
    Hill G E, McGraw K J (2006) Bird coloration. Harvard University Press, United Kingdom.
    Jacquin L, Lenouvel P, Haussy C, Ducatez S, Gasparini J (2011) Melanin‐based coloration is related to parasite intensity and cellular immune response in an urban free living bird: the feral pigeon Columba livia. J Avian Biol 42:11-15.
    Jawor J M, Breitwisch R (2003) Melanin ornaments, honesty, and sexual selection. Auk 120:249-265.
    Kingma S A, Szentirmai I, Szekely T, Bokony V, Bleeker M, Liker A, Komdeur J (2008) Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits Remiz pendulinus. Behav Ecol Sociobiol 62:1277-1288.
    Latscha T (1990) Carotenoids-their nature and significance in animal feeds. F. Hoffmann-La Roche & Co. Ltd., Basle.
    Mayr E (1972) Sexual selection and natural selection. 87-104.
    McGraw K J (2006a) Mechanics of carotenoid-based coloration. In: Hill (ed) Bird coloration, 3rd edn. Harvard Univ. Press Cambridge MA.
    McGraw K J (2006b) Mechanics of melanin-based coloration. In: Hill and McGraw (ed) Bird Coloration. Vol. I. Mechanisms and Measurement, 3rd edn. Harvard University Press, Cambridge.
    McGraw K J, Mackillop E A, Dale J, Hauber M E (2002) Different colors reveal different information: how nutritional stress affects the expression of melanin-and structurally based ornamental plumage. J Exp Biol 205:3747-3755.
    Mennill D J, Doucet S M, Montgomerie R, Ratcliffe L M (2003) Achromatic color variation in black-capped chickadees, Poecile atricapilla: black and white signals of sex and rank. Behav Ecol Sociob 53:350-357.
    Neff B D, Pitcher T E (2005) Genetic quality and sexual selection: an integrated framework for good genes and compatible genes. Mol Ecol 14:19-38.
    Olson V A, Owens I P F (1998) Costly sexual signals: are carotenoids rare, risky or required? Trends Ecol Evol 13:510-514.
    Pike T W, Blount J D, Lindstrom J, Metcalfe N B (2007) Dietary carotenoid availability influences a male's ability to provide parental care. Behav Ecol 18:1100-1105.
    Pryke S R, Andersson S, Lawes M J (2001) Sexual selection of multiple handicaps in the red‐collared widowbird: female choice of tail length but not carotenoid display. Evolution 55:1452-1463.
    Reudink M W, Studds C E, Marra P P, Kurt Kyser T, Ratcliffe L M (2009) Plumage brightness predicts non breeding season territory quality in a long distance migratory songbird, the American redstart Setophaga ruticilla. J Avian Biol 40:34-41.
    Roulin A, Gasparini J, Bize P, Ritschard M, Richner H (2008) Melanin-based colorations signal strategies to cope with poor and rich environments. Behav Ecol Sociobiol 62:507-519.
    Senar J C, Figuerola J, Pascual J (2002) Brighter yellow blue tits make better parents. Proc R Soc Lond B Biol Sci 269:257-261.
    Stevens M (2013) Sensory ecology, behaviour, and evolution. OUP Oxford, United kingdom.
    Tarof S A, Dunn P O, Whittingham L A (2005) Dual functions of a melanin-based ornament in the common yellowthroat. Proc R Soc Lond B Biol Sci 272:1121-1127.
    Walsh N, Dale J, McGraw K, Pointer M, Mundy N (2012) Candidate genes for carotenoid coloration in vertebrates and their expression profiles in the carotenoid-containing plumage and bill of a wild bird. Proc R Soc Lond B Biol Sci 279:58-66.
    Chapter Two
    Alonso-Alvarez C, Doutrelant C, Sorci G (2004) Ultraviolet reflectance affects male-male interactions in the blue tit (Parus caeruleus ultramarinus). Behav Ecol 15:805-809.
    Amundsen T, Forsgren E (2001) Male mate choice selects for female coloration in a fish. P Natl Acad Sci USA 98:13155-13160.
    Andersson M B (1994) Sexual selection. Princeton University Press, NJ.
    Armenta J K, Dunn P O, Whittingham L A (2008) Effects of specimen age on plumage color. Auk 125:803-808.
    Balakrishnan P (2010) Reproductive biology of the square-tailed Black Bulbul Hypsipetes ganeesa in the Western Ghats, India. Indian Birds 5:134-138.
    Bennett A T, Cuthill I C, Partridge J C, Lunau K (1997) Ultraviolet plumage colors predict mate preferences in starlings. P Natl Acad Sci USA 94:8618-8621.
    Borgia G (1995) Complex male display and female choice in the spotted bowerbird: specialized functions for different bower decorations. Anim Behav 49:1291-1301.
    Bortolotti G R, Negro J J, Tella J L, Marchant T A, Bird D M (1996) Sexual dichromatism in birds independent of diet, parasites and androgens. P Roy Soc B 263:1171-1176.
    Bridge E S, Hylton J, Eaton M D, Gamble L, Schoech S J (2008) Cryptic plumage signaling in Aphelocoma scrub-jays. J Ornithol 149:123-130.
    Bright A, Waas J R (2002) Effects of bill pigmentation and UV reflectance during territory establishment in blackbirds. Anim Behav 64:207-213.
    Buckley P A (1989) Medelian genes. In: Cook and Buckley (ed) Avian genetics. A population and ecological approach, 3rd edn. Academic Press, London.
    Chaine A S, Lyon B E (2008) Adaptive plasticity in female mate choice dampens sexual selection on male ornaments in the lark bunting. Science 319:459-462.
    Chen D-M, Collins J S, Goldsmith T H (1984) The ultraviolet receptor of bird retinas. 225:337-340.
    Cuthill I, Bennett A, Partridge J, Maier E (1999) Plumage reflectance and the objective assessment of avian sexual dichromatism. Am Nat 153:183-200.
    Dale J (2006) Intraspecific variation in coloration. In: Hill and McGraw (ed) Bird coloration, function and evolution, 3rd edn. President and Fellows of Harvard College, USA.
    Darwin C (1872) The origin of species by means of natural selection: Or, the preservation of favoured races in the struggle for life and the descent of man and selection in relation to sex. Modern Library,
    Delhey K, Peters A (2008) Quantifying variability of avian colours: are signalling traits more variable? PLOS ONE 3:e1689.
    Doucet S M, Hill G E (2009) Do museum specimens accurately represent wild birds? A case study of carotenoid, melanin, and structural colours in long‐tailed manakins Chiroxiphia linearis. J Avian Biol 40:146-156.
    Doucet S M, Montgomerie R (2003) Multiple sexual ornaments in satin bowerbirds: ultraviolet plumage and bowers signal different aspects of male quality. Behav Ecol 14:503-509.
    Dunn P O, Garvin J C, Whittingham L A, Freeman‐Gallant C R, Hasselquist D (2010) Carotenoid and melanin‐based ornaments signal similar aspects of male quality in two populations of the common yellowthroat. Funct Ecol 24:149-158.
    Eaton M D (2005) Human vision fails to distinguish widespread sexual dichromatism among sexually monochromatic birds. P Natl Acad Sci USA 102:10942-10946.
    Edward D A, Chapman T (2011) The evolution and significance of male mate choice. Trends Ecol Evol 26:647-654.
    Endler J, Mielke JR P (2005) Comparing entire colour patterns as birds see them. Biol J Linn Soc 86:405-431.
    Fishpool L D C, Tobias J A (2005) Family Pycnonotidae (Bulbuls). In: Del Hoyo, Elliot and Sargatal (ed) Handbook of the Birds of the World, Vol 10, 3rd edn. Lynx Editions, Barcelona.
    Fox D L (1976) Animal biochromes and structural colours: physical, chemical, distributional & physiological features of coloured bodies in the animal world. Univ of California Press, California.
    Fridolfsson A K, Ellegren H (1999) A simple and universal method for molecular sexing of non-ratite birds. J Avian Biol 30:116-121.
    Gemmell N, Akiyama S (1996) An efficient method for the extraction of DNA from vertebrate tissues. Trends Genet 12:338-339.
    Goldsmith T H (1990) Optimization, constraint, and history in the evolution of eyes. Q Rev Biol 65:281-322.
    Gray D A (1996) Carotenoids and sexual dichromatism in North American passerine birds. Am Nat 148:453-480.
    Griggio M, Valera F, Casas A, Pilastro A (2005) Males prefer ornamented females: a field experiment of male choice in the rock sparrow. Anim Behav 69:1243-1250.
    Hill G E (1992) Proximate basis of variation in carotenoid pigmentation in male house finches. Auk 109:U1-12.
    Hill G E (1996) Redness as a measure of the production cost of ornamental coloration. Ethol Ecol Evol 8:157-175.
    Igic B, Leuschner N, Parker K A, Ismar S M H, Gill B J, Lovegrove T G, Millar C D, Hauber M E (2010) Size dimorphism and avian perceived sexual dichromatism in a New Zealand endemic bird, the whitehead Mohoua albicilla. J Morphol 271:697-704.
    Johnstone R A (1994) Honest signalling, perceptual error and the evolution of'all-or-nothing'displays. P Roy Soc B 256:169-175.
    Jones K M, Monaghan P, Nager R G (2001) Male mate choice and female fecundity in zebra finches. Anim Behav 62:1021-1026.
    Kingma S A, Szentirmai I, Szekely T, Bokony V, Bleeker M, Liker A, Komdeur J (2008) Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits Remiz pendulinus. Behav Ecol Sociobiol 62:1277-1288.
    Kokko H, Johnstone R A (2002) Why is mutual mate choice not the norm? Operational sex ratios, sex roles and the evolution of sexually dimorphic and monomorphic signalling. Philost T Roy Soc B 357:319-330.
    Lessells C, Boag P T (1987) Unrepeatable repeatabilities: a common mistake. Auk 104:116-121.
    Mays Jr H L, McGraw K J, Ritchison G, Cooper S, Rush V, Parker R S (2004) Sexual dichromatism in the yellow breasted chat Icteria virens: spectrophotometric analysis and biochemical basis. J Avian Biol 35:125-134.
    McGraw K (2006a) Mechanics of carotenoid-based coloration. In: Hill and McGray (ed) Bird Coloration: Function and evolution, 3rd edn. Harvard University Press, USA.
    McGraw K J (2006b) Mechanics of melanin-based coloration. In: Hill and McGray (ed) Bird Coloration: Function and evolution, 3rd edn. Harvard University Press, USA.
    McNett G D, Marchetti K, Zink R (2005) Ultraviolet degradation in carotenoid patches: live versus museum specimens of wood warblers (Parulidae). Auk 122:793-802.
    Moller A, Pomiankowski A (1993) Why have birds got multiple sexual ornaments? Behav Ecol Sociobio 32:167-176.
    Montgomerie R (2006) Analyzing colors. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard University Press, USA.
    Neitz J, Jacobs G H (1986) Polymorphism of the long-wavelength cone in normal human colour vision. Nature 323:623 - 625.
    Nolan P M, Hill G E, Stoehr A M (1998) Sex, size, and plumage redness predict house finch survival in an epidemic. P Roy Soc B 265:961-965.
    Owens I P F, Hartley I R (1998) Sexual dimorphism in birds: why are there so many different forms of dimorphism? P Roy Soc B 265:397-407.
    Pohland G, Mullen P (2006) Preservation agents influence UV-coloration of plumage in museum bird skins. J Ornithol 147:464-467.
    Préault M, Deregnaucourt S, Sorci G, Faivre B (2002) Does beak coloration of male blackbirds play a role in intra and/or intersexual selection? Behav Process 58:91-96.
    Seddon N, Botero C A, Tobias J A, Dunn P O, MacGregor H E, Rubenstein D R, Uy J A C, Weir J T, Whittingham L A, Safran R J (2013) Sexual selection accelerates signal evolution during speciation in birds. P Roy Soc B 280:20131065.
    Siefferman L, Hill G E (2005) UV-blue structural coloration and competition for nestboxes in male eastern bluebirds. Anim Behav 69:67-72.
    Stoddard M C, Prum R O (2008) Evolution of avian plumage color in a tetrahedral color space: a phylogenetic analysis of new world buntings. Am Nat 171:755-776.
    Stoddard M C, Prum R O (2011) How colorful are birds? Evolution of the avian plumage color gamut. Behav Ecol 22:1042-1052.
    Tarof S A, Dunn P O, Whittingham L A (2005) Dual functions of a melanin-based ornament in the common yellowthroat. P Roy Soc B 272:1121-1127.
    Thompson C W, Hillgarth N, Leu M, McClure H E (1997) High parasite load in house finches (Carpodacus mexicanus) is correlated with reduced expression of a sexually selected trait. Am Nat 149:270-294.
    Vorobyev M, Osorio D (1998) Receptor noise as a determinant of colour thresholds. P Roy Soc B 265:351-358.
    Vorobyev M, Osorio D, Bennett A, Marshall N, Cuthill I (1998) Tetrachromacy, oil droplets and bird plumage colours. J Comp Phys A 183:621-633.
    Walker L K, Stevens M, Karadaş F, Kilner R M, Ewen J G (2013) A window on the past: male ornamental plumage reveals the quality of their early-life environment. P Roy Soc B 280:2012-2852.
    Chapter Three
    Alan R R, McWilliams S R, McGraw K J (2013) The importance of antioxidants for avian fruit selection during autumn migration. Wilson J Ornithol 125:513-525.
    Alonso-Alvarez C, Bertrand S, Devevey G, Gaillard M, Prost J, Faivre B, Sorci G (2004) An experimental test of the dose-dependent effect of carotenoids and immune activation on sexual signals and antioxidant activity. Am Nat 164:651-659.
    Ardia D R, Broughton D R, Gleicher M J (2010) Short-term exposure to testosterone propionate leads to rapid bill color and dominance changes in zebra finches. Horm Behav 58:526-532.
    Barbosa A, Palacios M J, Negro J J, Cuervo J (2013) Plasma carotenoid depletion during fasting in moulting penguins. J Ornithol 154:559-562.
    Bertrand S, Alonso-Alvarez C, Devevey G, Faivre B, Prost J, Sorci G (2006) Carotenoids modulate the trade-off between egg production and resistance to oxidative stress in zebra finches. Oecologia 147:576-584.
    Biard C, Surai P F, Moller A P (2005) Effects of carotenoid availability during laying on reproduction in the blue tit. Oecologia 144:32-44.
    Blount J D, Metcalfe N B, Birkhead T R, Surai P F (2003) Carotenoid modulation of immune function and sexual attractiveness in zebra finches. Science 300:125-127.
    Casagrande S, Pinxten R, Zaid E, Eens M (2015) Birds receiving extra carotenoids keep singing during the sickness phase induced by inflammation. Behav Ecol Sociobiol 69:1029-1037.
    Cherel Y, Leloup J, Le Maho Y (1988) Fasting in king penguin. II. Hormonal and metabolic changes during molt. Am J Physiol 254:R178-R14.
    Dale J (2006) Intraspecific variation in coloration. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard University Press, United Kingdom.
    Del Val E, Negro J J, Garrido-Fernández J, Jarén M, Borràs A, Cabrera J, Senar J C (2014) Seasonal variation of red carotenoid pigments in plasma of wild Crossbill males Loxia curvirostra. J Ornithol 155:211-218.
    Del Val E, Negro J J, Senar J C (2013) Seasonal variation of circulating plasma carotenoids in Mediterranean Great Tit males Parus major. Bird Study 60:459-465.
    Dey C J, Valcu M, Kempenaers B, Dale J (2015) Carotenoid‐based bill coloration functions as a social, not sexual, signal in songbirds (Aves: Passeriformes). J Evol Biol 28:250-258.
    Faivre B, Gregoire A, Preault M, Cezilly F, Sorci G (2003) Immune activation rapidly mirrored in a secondary sexual trait. Science 300:103.
    Fox H M, Vevers G (1960) The nature of animal colours. Macmillan, NY USA.
    Gross W, Siegel H (1983) Evaluation of the heterophil/lymphocyte ratio as a measure of stress in chickens. Avian Dis 27:972-979.
    Jenni L, Winkler R (1994) Moult and ageing of European passerines. Academic Pr, London.
    Klaassen M (1995) Moult and basal metabolic costs in males of two subspecies of stonechats: the European Saxicola torquata rubicula and the East African S. t. axillaris. Oecologia 104:424-432.
    Klomp H (1970) The determination of clutch-size in birds: a review. Ardea 58:1-142.
    Kuenzel W (2003) Neurobiology of molt in avian species. Poult Sci 82:981-991.
    Latscha T (1990) Carotenoids-their nature and significance in animal feeds. F. Hoffmann-La Roche & Co. Ltd., Basle.
    Lindstrom A, Visser G H, Daan S (1993) The energetic cost of feather synthesis is proportional to basal metabolic rate. Physiol Zool 66:490-510.
    Lochmiller R L, Deerenberg C (2000) Trade‐offs in evolutionary immunology: just what is the cost of immunity? Oikos 88:87-98.
    Martin L B (2005) Trade-offs between molt and immune activity in two populations of house sparrows (Passer domesticus). Can J Zool 83:780-787.
    McGraw K, Adkins-Regan E, Parker R (2005) Maternally derived carotenoid pigments affect offspring survival, sex ratio, and sexual attractiveness in a colorful songbird. Naturwissenschaften 92:375-380.
    McGraw K, Nolan P, Crino O (2006) Carotenoid accumulation strategies for becoming a colourful house finch: analyses of plasma and liver pigments in wild moulting birds. Funct Ecol 20:678-688.
    Metzger B J, Bairlein F (2011) Fat stores in a migratory bird: a reservoir of carotenoid pigments for times of need? J Comp Physiol B 181:269-275.
    Ninni P, De Lope F, Saino N, Haussy C, Moller A P (2004) Antioxidants and condition-dependence of arrival date in a migratory passerine. Oikos 105:55-64.
    Nordeide J, Mohus A, Nicolaisen O, Volden R, Egeland E (2008) Offspring or ornaments? Is carotenoid based ornamentation in female Arctic charr, Salvelinus alpinus (L.), condition dependent and traded off against offspring? Ecol Freshw Fish 17:328-339.
    Olson V A, Owens I P F (1998) Costly sexual signals: are carotenoids rare, risky or required? Trends Ecol Evol 13:510-514.
    Pap P L, Vagasi C I, Czirjak G A, Barta Z (2008) Diet quality affects postnuptial molting and feather quality of the house sparrow (Passer domesticus): interaction with humoral immune function? Can J Zool 86:834-842.
    Perrins C (1970) The timing of birds' breeding seasons. Ibis 112:242-255.
    Rehder N, Bird D, Lague P (1986) Variations in plasma corticosterone, estrone, estradiol-17β, and progesterone concentrations with forced renesting, molt, and body weight of captive female American kestrels. Gen Comp Endocrinol 62:386-393.
    Sanz J J, Moreno J, Merino S, Tomas G (2004) A trade-off between two resource-demanding functions: post-nuptial moult and immunity during reproduction in male pied flycatchers. J Anim Ecol 73:441-447.
    Senar J C (2006) Color displays as intrasexual signals of aggression and dominance. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard University, United Kingdom.
    Serra L, Griggio M, Licheri D, Pilastro A (2007) Moult speed constrains the expression of a carotenoid based sexual ornament. J Evol Biol 20:2028-2034.
    Smits J, Bortolotti G R, Tella J L (1999) Simplifying the phytohaemagglutinin skin‐testing technique in studies of avian immunocompetence. Func Ecol 13:567-572.
    Spencer K, Buchanan K, Goldsmith A, Catchpole C (2003) Song as an honest signal of developmental stress in the zebra finch (Taeniopygia guttata). Horm Behav 44:132-139.
    Stevens M (2013) Sensory ecology, behaviour, and evolution. OUP Oxford, United kingdom.
    Surai P F (2002) Natural antioxidants in avian nutrition and reproduction. Nottingham University Press Nottingham, Nottingham.
    Vaugoyeau M, Decencière B, Perret S, Karadas F, Meylan S, Biard C (2015) Is oxidative status influenced by dietary carotenoid and physical activity after moult in the great tit (Parus major)? J Exp Bio jeb. 111039.
    Vleck C M, Vertalino N, Vleck D, Bucher T L (2000) Stress, corticosterone, and heterophil to lymphocyte ratios in free-living Adelie penguins. Condor 102:392-400.
    Zera A J, Harshman L G (2001) The physiology of life history trade-offs in animals. Annu Rev Ecol Syst 32:95-126.
    Chapter Four
    Andersson M, Simmons L W (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296-302.
    Badyaev A V, Hill G E (2000) Evolution of sexual dichromatism: contribution of carotenoid‐versus melanin‐based coloration. Biol J Linn Soc 69:153-172.
    Benedetto J-P, Ortonne J-P, Voulot C, Khatchadourian C, Prota G, Thivolet J (1982) Role of thiol compounds in mammalian melanin pigmentation. II. Glutathione and related enzymatic activities. J Invest Dermatol 79:422-424.
    Bize P, Gasparini J, Klopfenstein A, Altwegg R, Roulin A (2006) Melanin-based coloration is a nondirectionally selected sex-specifec signal of offspring development in the Alpine swift. Evolution 60:2370-2380.
    Davis A, Maney D, Maerz J (2008) The use of leukocyte profiles to measure stress in vertebrates: a review for ecologists. Funct Ecol 22:760-772.
    Ducrest A-L, Keller L, Roulin A (2008) Pleiotropy in the melanocortin system, coloration and behavioural syndromes. Trends Ecol Evol 23:502-510.
    Fargallo J A, Laaksonen T, Korpimäki E, Wakamatsu K (2007) A melanin-based trait reflects environmental growth conditions of nestling male Eurasian kestrels. Evol Ecol 21:157-171.
    Finkel T, Holbrook N J (2000) Oxidants, oxidative stress and the biology of ageing. Nature 408:239-247.
    Fridolfsson A-K, Ellegren H (1999) A simple and universal method for molecular sexing of non-ratite birds. J Avian Biol 116-121.
    Galván I, Alonso-Alvarez C (2008) An intracellular antioxidant determines the expression of a melanin-based signal in a bird. PLoS One 3:e3335.
    Galván I, Alonso-Alvarez C (2009) The expression of melanin-based plumage is separately modulated by exogenous oxidative stress and a melanocortin. Proc R Soc B 276:3089-3097.
    Gemmell N J, Akiyama S (1996) An efficient method for the extraction of DNA from vertebrate tissues. Trends Genet 12:338-339.
    Griffith S C, Parker T H, Olson V A (2006) Melanin-versus carotenoid-based sexual signals: is the difference really so black and red? Anim Behav 71:749-763.
    Gross W, Siegel H (1983) Evaluation of the heterophil/lymphocyte ratio as a measure of stress in chickens. Avian Dis 27:972-979.
    Hõrak P, Sild E, Soomets U, Sepp T, Kilk K (2010) Oxidative stress and information content of black and yellow plumage coloration: an experiment with greenfinches. J Exp Bio 213:2225-2233.
    Hill G E, Brawner W R (1998) Melanin–based plumage coloration in the house finch is unaffected by coccidial infection. Proc R Soc B 265:1105-1109.
    Hill G E, McGraw K J (2006) Bird coloration I: mechanisms and measurements. Harvard University Press, Cambridge.
    Hung H-Y, Yeung C K L, Li S-H (2015) Sexual dichromatism invisible to the human eye and the quality of museum specimens in the Himalayan black bulbul (Hypsipetes leucocephalus niggerimus, Pycnonotidae). Zool Stud in revision.
    Jacquin L, Lenouvel P, Haussy C, Ducatez S, Gasparini J (2011) Melanin‐based coloration is related to parasite intensity and cellular immune response in an urban free living bird: the feral pigeon Columba livia. J Avian Biol 42:11-15.
    Jawor J M, Breitwisch R (2003) Melanin ornaments, honesty, and sexual selection. Auk 120:249-265.
    Maxwell M (1993) Avian blood leucocyte responses to stress. World Poult Sci J 49:34-43.
    McFarlane J M, Curtis S E (1989) Multiple concurrent stressors in chicks. 3. Effects on plasma corticosterone and the heterophil: lymphocyte ratio. Poult Sci 68:522-527.
    McGraw K, Safran R, Wakamatsu K (2005) How feather colour reflects its melanin content. Func Ecol 19:816-821.
    McGraw K J (2006) Mechanics of melanin-based coloration. In: Hill and McGraw (ed) Bird Coloration. Vol. I. Mechanisms and Measurement, 3rd edn. Harvard University Press, Cambridge.
    Meister A (1983) Selective modification of glutathione metabolism. Science 220:472-477.
    Monaghan P, Metcalfe N B, Torres R (2009) Oxidative stress as a mediator of life history trade‐offs: mechanisms, measurements and interpretation. Ecol Lett 12:75-92.
    Patel R, McIntosh L, McLaughlin J, Brooke S, Nimon V, Sapolsky R (2002) Disruptive effects of glucocorticoids on glutathione peroxidase biochemistry in hippocampal cultures. J Neurochem 82:118-125.
    Roulin A, Almasi B, Rossi-Pedruzzi A, Ducrest A-L, Wakamatsu K, Miksik I, Blount J D, Jenni-Eiermann S, Jenni L (2008) Corticosterone mediates the condition-dependent component of melanin-based coloration. Anim Behav 75:1351-1358.
    Vleck C M, Vertalino N, Vleck D, Bucher T L (2000) Stress, corticosterone, and heterophil to lymphocyte ratios in free-living Adelie penguins. Condor 102:392-400.
    Chapter Five
    Andersson M, Simmons L W (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296-302.
    Andersson M B (1994) Sexual selection. Princeton University Press, NJ USA.
    Ardia D R, Broughton D R, Gleicher M J (2010) Short-term exposure to testosterone propionate leads to rapid bill color and dominance changes in zebra finches. Horm Behav 58:526-532.
    Blount J D, Matheson S M (2006) Effects of carotenoid supply on escape flight responses in zebra finches, Taeniopygia guttata. Anim Behav 72:595-601.
    Blount J D, Metcalfe N B, Birkhead T R, Surai P F (2003) Carotenoid modulation of immune function and sexual attractiveness in zebra finches. Science 300:125-127.
    Borgia G (1995) Complex male display and female choice in the spotted bowerbird: specialized functions for different bower decorations. Anim Behav 49:1291-1301.
    Brush A H (1990) Metabolism of carotenoid pigments in birds. FASEB J 4:2969-2977.
    Burley N, Coopersmith C B (1987) Bill color preferences of zebra finches. Ethology 76:133-151.
    Casagrande S, Csermely D, Pini E, Bertacche V, Tagliavini J (2006) Skin carotenoid concentration correlates with male hunting skill and territory quality in the kestrel Falco tinnunculus. J Avian Biol 37:190-196.
    Chaine A S, Lyon B E (2008) Adaptive plasticity in female mate choice dampens sexual selection on male ornaments in the lark bunting. Science 319:459-462.
    Dey C J, Valcu M, Kempenaers B, Dale J (2015) Carotenoid‐based bill coloration functions as a social, not sexual, signal in songbirds (Aves: Passeriformes). J Evol Biol 28:250-258.
    Faivre B, Gregoire A, Preault M, Cezilly F, Sorci G (2003) Immune activation rapidly mirrored in a secondary sexual trait. Science 300:103.
    Fridolfsson A K, Ellegren H (1999) A simple and universal method for molecular sexing of non-ratite birds. J Avian Biol 30:116-121.
    Gemmell N J, Akiyama S (1996) An efficient method for the extraction of DNA from vertebrate tissues. Trends Genet 12:338-339.
    Griggio M, Serra L, Licheri D, Monti A, Pilastro A (2007) Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal. Behav Ecol Sociobiol 61:423-433.
    Hill G E (2006) Female mate choice for ornamental coloration. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard Universtiy Press, United Kingkom.
    Horak P, Ots I, Vellau H, Spottiswoode C, Pape Moller A (2001) Carotenoid-based plumage coloration reflects hemoparasite infection and local survival in breeding great tits. Oecologia 126:166-173.
    Hung H-Y, Yeung C K L, Li S-H (2015a) Sexual dichromatism invisible to the human eye and the quality of museum specimens in the Himalayan black bulbul (Hypsipetes leucocephalus nigerrimus, Pycnonotidae). Zool Stud In revision:
    Hung H-Y, Yeung C K L, Li S-H (2015b) Sexual dichromatism invisible to the human eye and the quality of museum specimens in the Himalayan black bulbul (Hypsipetes leucocephalus niggerimus, Pycnonotidae). Zool Stud in revision.
    Inouye C Y, Hill G E, Stradi R D, Montgomerie R, Bosque C (2001) Carotenoid pigments in male house finch plumage in relation to age, subspecies, and ornamental coloration. Auk 118:900-915.
    Jawor J M, Gray N, Beall S M, Breitwisch R (2004) Multiple ornaments correlate with aspects of condition and behaviour in female northern cardinals, Cardinalis cardinalis. Anim Behav 67:875-882.
    Johnstone R A (1994) Honest signalling, perceptual error and the evolution of'all-or-nothing'displays. P Roy Soc B 256:169-175.
    McGraw K J (2006) Mechanics of carotenoid-based coloration. In: Hill (ed) Bird coloration, 3rd edn. Harvard Univ. Press Cambridge MA.
    Moller A, Pomiankowski A (1993) Why have birds got multiple sexual ornaments? Behav Ecol Sociobio 32:167-176.
    Nolan P M, Dobson F S, Dresp B, Jouventin P (2006) Immunocompetence is signalled by ornamental colour in king penguins, Aptenodytes patagonicus. Evol Ecol Res 8:1325-1332.
    Olson V A, Owens I P F (1998) Costly sexual signals: are carotenoids rare, risky or required? Trends Ecol Evol 13:510-514.
    Pryke S R, Andersson S (2003) Carotenoid-based status signalling in red-shouldered widowbirds (Euplectes axillaris): epaulet size and redness affect captive and territorial competition. Behav Ecol Soci 53:393-401.
    Pryke S R, Andersson S, Lawes M J, Piper S E (2002) Carotenoid status signaling in captive and wild red-collared widowbirds: independent effects of badge size and color. 13:622-631.
    Pryke S R, Lawes M J, Andersson S (2001) Agonistic carotenoid signalling in male red-collared widowbirds: aggression related to the colour signal of both the territory owner and model intruder. Anim Behav 62:695-704.
    Qvarnström A, Pärt T, Sheldon B C (2000) Adaptive plasticity in mate preference linked to differences in reproductive effort. 405:344-347.
    Saks L, Ots I, Horak P (2003) Carotenoid-based plumage coloration of male greenfinches reflects health and immunocompetence. Oecologia 134:301-307.
    Senar J, Figuerola J, Pascual J (2002) Brighter yellow blue tits make better parents. Proc R Soc Lond B Biol Sci 269:257-261.
    Senar J C (2006) Color displays as intrasexual signals of aggression and dominance. In: Hill and McGraw (ed) Bird coloration, 3rd edn. Harvard University, United Kingdom.
    Severinghaus L L, Ding T-s, Fang W-H, Lin W-H, Tsa i M-C, Yen C-W (2012) The avifauna of Taiwan. Forest Bureau, Taipei.
    Stevens M (2013) Sensory ecology, behaviour, and evolution. OUP Oxford, United kingdom.
    Svensson P A, Wong B (2011) Carotenoid-based signals in behavioural ecology: a review. Behaviour 148:131-189.
    Thomas J, Gausman H (1977) Leaf reflectance vs. leaf chlorophyll and carotenoid concentrations for eight crops. Agron J 69:799-802.
    Viera V M, Nolan P M, Côté S D, Jouventin P, Groscolas R (2008) Is territory defence related to plumage ornaments in the king penguin Aptenodytes patagonicus? Ethology 114:146-153.
    West-Eberhard M J (1983) Sexual selection, social competition, and speciation. Q Rev Biol 58:155-183.
    Chapter Six
    Andersson M (1994) Sexual Selection. Princeton University Press, Princeton.
    Arch V S, Grafe T U, Narins P M (2008) Ultrasonic signalling by a Bornean frog. 4:19-22.
    Blount J D, Metcalfe N B, Birkhead T R, Surai P F (2003) Carotenoid modulation of immune function and sexual attractiveness in zebra finches. Science 300:125-127.
    Borgia G (1995) Complex male display and female choice in the spotted bowerbird: specialized functions for different bower decorations. Anim Behav 49:1291-1301.
    Chaine A S, Lyon B E (2008) Adaptive plasticity in female mate choice dampens sexual selection on male ornaments in the lark bunting. Science 319:459-462.
    Dale J (2006) Intraspecific Variation in Coloration. In: Hill and McGraw (ed) Bird coloration: function and evolution 3rd edn. Harvard University Press, United Kingdom.
    Delhey K, Peters A (2008) Quantifying variability of avian colours: are signalling traits more variable? PLOS ONE 3:e1689.
    Dey C J, Valcu M, Kempenaers B, Dale J (2015) Carotenoid‐based bill coloration functions as a social, not sexual, signal in songbirds (Aves: Passeriformes). J Evol Biol 28:250-258.
    Dunn P O, Armenta J K, Whittingham L A (2015) Natural and sexual selection act on different axes of variation in avian plumage color. Sci Adv 1:e1400155.
    Faivre B, Gregoire A, Preault M, Cezilly F, Sorci G (2003a) Immune activation rapidly mirrored in a secondary sexual trait. Science 300:103.
    Faivre B, Préault M, Salvadori F, Théry M, Gaillard M, Cézilly F (2003b) Bill colour and immunocompetence in the European blackbird. Anim Behav 65:1125-1131.
    Gil D, Gahr M (2002) The honesty of bird song: multiple constraints for multiple traits. Trends Ecol Evol 17:133-141.
    Hunt S, Cuthill I C, Bennett A T, Griffiths R (1999) Preferences for ultraviolet partners in the blue tit. Anim Behav 58:809-815.
    Jawor J M, Breitwisch R (2003) Melanin ornaments, honesty, and sexual selection. Auk 120:249-265.
    Jawor J M, Breitwisch R (2004) Multiple ornaments in male northern cardinals, Cardinalis cardinalis, as indicators of condition. Ehtology 110:113-126.
    Jawor J M, Linville S U, Beall S M, Breitwisch R (2003) Assortative mating by multiple ornaments in northern cardinals (Cardinalis cardinalis). Behav Ecol 14:515-520.
    Johnstone R A (1994) Honest signalling, perceptual error and the evolution of'all-or-nothing'displays. P Roy Soc B 256:169-175.
    Jouventin P, Nolan P M, Örnborg J, Dobson F S (2005) Ultraviolet beak spots in king and emperor penguins. Condor 107:144-150.
    Li J, Zhang Y, Zhang Z (2011) High frequency components in avian vocalizations. Chinese birds 2:125-131.
    Moller A, Pomiankowski A (1993) Why have birds got multiple sexual ornaments? Behav Ecol Sociobio 32:167-176.
    Mougeot F, Pérez‐Rodríguez L, Sumozas N, Terraube J (2009) Parasites, condition, immune responsiveness and carotenoid‐based ornamentation in male red‐legged partridge Alectoris rufa. J Avian Biol 40:67-74.
    Nolan P M, Stephen Dobson F, Nicolaus M, Karels T J, McGraw K J, Jouventin P (2010) Mutual mate choice for colorful traits in king penguins. Ethology 116:635-644.
    Norris K (1990) Female choice and the quality of parental care in the great tit Parus major. Behav Ecol Sociobiol 27:275-281.
    Pérez-Rodríguez L, Viñuela J (2008) Carotenoid-based bill and eye ring coloration as honest signals of condition: an experimental test in the red-legged partridge (Alectoris rufa). Naturwissenschaften 95:821-830.
    Peters A, Denk A, Delhey K, Kempenaers B (2004) Carotenoid‐based bill colour as an indicator of immunocompetence and sperm performance in male mallards. J Evol Biol 17:1111-1120.
    Pytte C L, Ficken M S, Moiseff A (2004) Ultrasonic singing by the blue-throated hummingbird: a comparison between production and perception. J Comp Physiol A 190:665-673.
    Qvarnström A, Pärt T, Sheldon B C (2000) Adaptive plasticity in mate preference linked to differences in reproductive effort. 405:344-347.
    Rosenthal M F, Murphy T G, Darling N, Tarvin K A (2012) Ornamental bill color rapidly signals changing condition. J Avian Biol 43:553-564.
    Roulin A, Gasparini J, Bize P, Ritschard M, Richner H (2008) Melanin-based colorations signal strategies to cope with poor and rich environments. Behav Ecol Sociobiol 62:507-519.
    Shutler D, Weatherhead P J (1990) Targets of sexual selection: song and plumage of wood warblers. 1967-1977.
    Viera V M, Nolan P M, Côté S D, Jouventin P, Groscolas R (2008) Is territory defence related to plumage ornaments in the king penguin Aptenodytes patagonicus? Ethology 114:146-153.

    下載圖示
    QR CODE