臺灣博碩士論文加值系統

根據本實驗室夜間野外調查，雄褐樹蛙於繁殖季會呈現明亮金黃體色，此與其性腺指數、血液睪固酮含量與繁殖季節(四月到六月)相符合。但對於雄褐樹蛙體色的改變、體內荷爾蒙與繁殖行為的關聯性並不明瞭。由背景適應實驗經Photoshop軟體量化體色之hue, saturation及 brightness，且利用USB2000 反射光譜儀測得繁殖季節中雄蛙金黃色的反射光譜圖為610 nm，結果得知於黃色背景下雄蛙無法改變為明亮金黃體色達到保護色作用，此現象為動態性別二原色 (dynamic sexual dichromatism)。藉由組織切片與共軛焦顯微鏡影像顯示褐樹蛙皮膚的色素細胞位於疏鬆層，當黃色色素擴散並移到疏鬆層上層覆蓋聚集的黑色色素，導致雄蛙呈金黃體色。在離體皮膚實驗中，睪固酮 (Testosterone, T)及泌乳激素 (Prolactin, PRL)會刺激黃色色素擴散，退黑激素 (Melatonin, MEL)則使黑色色素聚集，但雌二醇 (Estradiol, E2)則無影響。當PRL與Ｔ同時處理時，黃色色素明顯的擴散。T+MEL, PRL+MEL及T+PRL+MEL等複合激素處理，發現黃色色素大量擴散往疏鬆層上層移動，並覆蓋聚集在下層的黑色色素，因此使皮膚顏色變黃。另外，在黑色背景下雄褐樹蛙呈現深棕色，腦下垂體的中的腦源性神經營養因子 (Brain-derived neurotrophic factor, BDNF)與腦下垂體的垂體腺苷酸環化酶激活肽 (Pituitary adenylate cyclase-activating polypeptide, PACAP) 較白色背景下呈淺棕色時表現量高，然而在任何背景條件下金黃體色時，褐樹蛙的BDNF及 PACAP表現量最低，意味著可能藉由BDNF 與 PACAP表現量的下降，因而減少黑色素細胞刺激素 (α-melanocyte stimulating hormone, α-MSH)的分泌，導致黑色色素聚集，使體色較為明亮，結果顯示體內不同激素於繁殖季中，可能扮演著調節褐樹蛙金黃體色的相關因子。進一步分析褐樹蛙視網膜中的視桿狀細胞(rod cell)及視錐狀細胞(cone cell)，發現cone cell有不同顏色的彩色油滴包括黃，橘，紫，綠，紅和藍色，以488, 543及405 nm激發光，找出其中三種不同吸收波長範圍的彩色油滴分別為498-632 nm (黃色、綠色及橘色)，582-632 nm (橘色) 和458-532 nm (綠色及藍色)，意味褐樹蛙擁有彩色視覺。並進一步進行喜好行為實驗探討雌蛙是否利用彩色視覺區分種間的金黃體色雄蛙，結果指出雌褐樹蛙明顯偏好金黃體色雄蛙，且不受明亮的螢幕吸引，另外發現雄蛙鳴叫聲會促進雄蛙對雌蛙的吸引力，這些結果與野外繁殖季時，雄蛙會停留在棲息地的大石頭上鳴叫，並呈現金黃體色達到吸引雌蛙靠近相符合，綜觀褐樹蛙為兩性大小異形(sexual size dimorphism)，動態性別二元色及雄褐樹蛙呈現明亮金黃體色為重要的生殖指標。

According to our field observations at night, the male Buergeria robusta in the breeding season showed brilliant yellow body color highly correlate (high somatic gonad index and plasma testosterone levels) to breeding season (from April to June) but not found in female. However, the relationship among body color change, hormone and breeding behavior is not clear yet. The background adaptation experiments showed that body color of male frog did not change to brilliant yellow in yellow background to achieve camouflage by quantification of body coloration (hue, saturation and brightness) with Photoshop software. Using USB2000 spectrometer measured the refractory spectrum of dorsal skin exhibited a dramatic peak at yellow spectrum (610 nm) in brilliant yellow skin of male frog during the breeding season. This observation demonstrates that B. robusta is a dynamic sexual dichromatism. The hematoxylin and eosin staining and confocal imaging were employed to detect the movement of pigments, chromatophores existed in stratum spongiosum layer of skin and yellow pigments in upper layer horizontally extend to cover the black pigment in low layer resulting in brilliant yellow skin. In isolated frog skin experiments, testosterone (T) and prolactin (PRL) were stimulated the dispersion of yellow pigments and the black pigments were aggregated by melatonin (MEL) treatment, but not estradiol (E2). In PRL+T treatment, yellow pigments were significantly dispersed. In T+MEL, PRL+MEL, and T+PRL+MEL treatments, yellow pigments were dispersed and the aggregations of black pigments were moved to the low layer of spongiosum in dorsal skin caused the increase of yellow coloration. In addition, brain-derived neurotrophic factor (BDNF) and pituitary adenylate cyclase-activating polypeptide (PACAP) expressions of pituitary from dark brown B. robusta in black background were higher than those of pale brown male frog in white background. Whereas BDNF and PACAP were the lowest expressions from brilliant yellow body color of male frog in any background condition, implying that lower expressions of PACAP and BDNF may reduce the α-melanocyte stimulating hormone (α-MSH) production resulted in the gathering of black pigment and consequently become brilliant yellow body color appearance. These results suggest that internal hormone may play a regulatory role in coloration of male B. robusta during the breeding season. Moreover, confocal microscope with excitation wavelength at 488, 543, and 405 nm to analyze the rod cell and cone cell of retina in B. robusta showed three different oil droplets in the absorption wavelength of 498-632 nm (yellow, green and orange), 582-632 nm (orange), and 458-532 nm (green and blue), respectively, revealing that B. robusta may have the color vision. Furthermore, preference behavior test was performed to measure whether female B. robusta may distinguish interspecies male frog with brilliant yellow body color by color vision. Female frog significantly preferred the male with brilliant yellow body color but not affected by the brightness screen alone. Additionally, the male calling could promote the attraction of male frog to female choice. These results correspond with wild field observation during the breeding season, the male frog changed body color to brilliant yellow stands on the rock at habitat area with calling to attract female frog. Taken altogether, brilliant yellow body color of male B. robusta is an important mating indicator for female seeking male by color vision during the breeding season.

Table contents IX
Figure contents X
Supplement contents XII
Abbreviations XIII
Introduction 1
Brown tree frog - Buergeria robusta 1
Sexual size dimorphism 2
Camouflage, sexual dichromatism and dynamic sexual dichromatism 3
Chromatophores 4
Body coloration is mediated by hormone 5
Regulatory factors of body coloration in pituitary gland. 7
a. Proopiomelanocortin (POMC), α-melanocyte stimulating hormone (α-MSH) and Melanotrope cell 7
b. Pituitary Adenylate Cyclase-Activating Polypeptide (PACAP) 8
c. Brain-Derived Neurotrophic Factor (BDNF) 9
Color vision 10
Sexual selection and mate choice of female 12
Motivation and Aims 15
Research design 17
(I) External factors and internal factors 17
(II) Animal Behavior 19
Materials and Methods 21
1、Buergeria robusta 21
2、Background adaptation experiments 21
2.1、Quantification of coloration 22
2.1.1 Adobes PhotoShops CS3 22
2.1.2 USB 2000 spectrometer 23
3、Morphology of chromatophore assay 23
3.1 Confocal microscope imaging 23
3.2 Hematoxylin and eosin staining 23
4、Measurement of blood Testosterone 24
5、Hormone treatment in vitro 25
5.1 Chemicals 25
5.2 Single hormone treatment and combinations treatment 25
6、Injection experiment in vivo 26
7、Total RNA isolation 26
8、Reverse transcription-polymerase chain reaction (RT-PCR) 26
9、Molecular cloning 27
10、Visual analysis 28
10.1 Hematoxylin and eosin staining 28
10.2 Colored oil droplets of cone cells assay 29
11、Animal behavior experiments 29
11.1 Installing the behavior arena 29
11.2 The protocol of preference test 30
11.3 Experiment 1 (exp. 1): hearing effect 32
12.4 Experiment 2 (exp. 2): olfaction effect 33
12.5 Experiment 3 (exp. 3): olfaction with hearing effect 34
12.6 Experiment 4 (exp. 4): visual effect. 35
12.6.1 Experiment 4-1 (exp. 4-1): effect of screen brightness. 35
12.6.2 Experiment 4-2 (exp. 4-2): effect of body coloration - first choice. 36
12.6.3 Experiment 4-3 (exp. 4-3): influence of body coloration- second choice. 37
12.7 Experiment 5 (exp. 5): visual with hearing experiments 38
12.7.1 Experiment 5-1 (exp. 5-1): effect of screen brightness with male calling 38
12.7.2 Experiment 5-2 (exp. 5-2): effect of body coloration with male calling-first choice. 39
12.7.3 Experiment 5-3 (exp. 5-3): effect of body coloration with male calling- second choice. 40
12.8 Experiment 6 (exp. 6): effect of visual with hearing and olfaction. 41
12.9 Experiment 7 (exp. 7): effect of male 42
12.10 Experiment 8 (exp. 8): effect of different species image experiments. 43
12.10.1 Experiment 8-1 (exp. 8-1) different species of image experiments with brown body color of male B. robusta and Bufo bankorensis. 43
12.10.2 Experiment 8-2 (exp. 8-2) different species of image experiments with brilliant yellow body color of male B. robusta and Bu. bankorensis. 44
Results 47
Field observations and background adaptation experiments (camouflage, sexual dichromatism and dynamic sexual dichromatism) 47
Quantification of body color 47
The morphology of chromatophore 48
Breeding season 49
Single and combination of hormone treatment in vitro 50
Injection experiment in vivo 51
Regulatory factors of body coloration in pituitary 51
Visual analysis and oil droplets of cone cells assay 52
Animal behavior experiments 53
Discussion 57
Breeding season 57
Dynamic sexual dichromatism 58
Sexual selection 59
The impact of sensation on breeding behavior 60
Color vision 61
Hormonal regulation of body color 63
Conclusion 67
Future works 69
References 71
Supplement 123

Aho, A. C., K. Donner, S. Helenius, L. O. Larsen and T. Reuter (1993). "Visual performance of the toad (Bufo bufo) at low light levels: retinal ganglion cell responses and prey-catching accuracy." J Comp Physiol A 172(6): 671-682.
Aho, A. C., K. Donner, C. Hyden, L. O. Larsen and T. Reuter (1988). "Low retinal noise in animals with low body temperature allows high visual sensitivity." Nature 334(6180): 348-350.
Amundsen, T. and E. Forsgren (2003). "Male preference for colourful females affected by male size in a marine fish." Behav Ecol Sociobiol 54(1): 55-64.
Andersson, M. (1994). Sexual selection, Princeton University Press.
Andersson, M. and Y. Iwasa (1996). "Sexual selection." Trends Ecol Evol 11(2): 53-58.
Arafa, S., M. Chouaibi, S. Sadok and A. El Abed (2012). "The influence of season on the gonad index and biochemical composition of the sea urchin Paracentrotus lividus from the Golf of Tunis." ScientificWorldJournal 2012: 815935.
Ardia, D. R., D. R. Broughton and M. J. Gleicher (2010). "Short-term exposure to testosterone propionate leads to rapid bill color and dominance changes in zebra finches." Horm Behav 58(3): 526-532.
Aspengren, S., D. Hedberg and M. Wallin (2006). "Studies of pigment transfer between Xenopus laevis melanophores and fibroblasts in vitro and in vivo." Pigment Cell Res 19(2): 136-145.
Aspengren, S., H. N. Sköld, G. Quiroga, L. Mårtensson and M. Wallin (2003). "Noradrenaline‐and Melatonin‐Mediated Regulation of Pigment Aggregation in Fish Melanophores." Pigment Cell Res 16(1): 59-64.
Badyaev, A. V. and G. E. Hill (2003). "Avian sexual dichromatism in relation to phylogeny and ecology." Annu Rev Ecol Evol 34: 27-49.
Bailes, H. J., S. R. Robinson, A. E. Trezise and S. P. Collin (2006). "Morphology, characterization, and distribution of retinal photoreceptors in the Australian lungfish Neoceratodus forsteri (Krefft, 1870)." J Comp Neurol 494(3): 381-397.
Beeching, S. C., S. H. Gross, H. S. Bretz and E. Hariatis (1998). "Sexual dichromatism in convict cichlids: the ethological significance of female ventral coloration." Anim Behav 56(4): 1021-1026.
Bell, R. C. and K. R. Zamudio (2012). "Sexual dichromatism in frogs: natural selection, sexual selection and unexpected diversity." Proc Biol Sci 279(1748): 4687-4693.
Bertrand, S., B. Faivre and G. Sorci (2006). "Do carotenoid-based sexual traits signal the availability of non-pigmentary antioxidants?" J Exp Biol 209(22): 4414-4419.
Bowmaker, J. K. (1977). "The visual pigments, oil droplets and spectral sensitivity of the pigeon." Vision Res 17(10): 1129-1138.
Bowmaker, J. K., E. R. Loew and M. Ott (2005). "The cone photoreceptors and visual pigments of chameleons." J Comp Physiol A Neuroethol Sens Neural Behav Physiol 191(10): 925-932.
Bradbury, J. W. and S. L. Vehrencamp (1998). "Principles of animal communication." Behav Ecol 12: 283-286.
Braekevelt, C. R. (1992). "Retinal photoreceptor fine structure in the red-backed salamander (Plethodon cinereus)." Histol Histopathol 7(3): 463-470.
Bramham, C. R. and E. Messaoudi (2005). "BDNF function in adult synaptic plasticity: the synaptic consolidation hypothesis." Prog Neurobiol 76(2): 99-125.
Briscoe, A. D. and L. Chittka (2001). "The evolution of color vision in insects." Annu Rev Entomol 46(1): 471-510.
Candolin, U. (2003). "The use of multiple cues in mate choice." BioRev 78(4): 575-595.
Cox, R. M., V. Zilberman and H. B. John-Alder (2008). "Testosterone stimulates the expression of a social color signal in Yarrow's Spiny Lizard, Sceloporus jarrovii." J Exp Zool A Ecol Genet Physiol 309(9): 505-514.
Cummings, M. E., X. E. Bernal, R. Reynaga, A. S. Rand and M. J. Ryan (2008). "Visual sensitivity to a conspicuous male cue varies by reproductive state in Physalaemus pustulosus females." J Exp Biol 211(Pt 8): 1203-1210.
Czeczuga, B. (1980). "Investigations on carotenoids in Amphibia—II. Carotenoids occurring in various parts of the body of certain species." Comp Bio Phy B 65(4): 623-630.
De Loof, A., J. Huybrechts, M. Geens, T. Vandersmissen, B. Boerjan and L. Schoofs (2010). "Sexual differentiation in adult insects: Male-specific cuticular yellowing in Schistocerca gregaria as a model for reevaluating some current (neuro)endocrine concepts." J Insect Physiol 56(8): 919-925.
de Luna, A. G., W. Hodl and A. Amezquita (2010). "Colour, size and movement as visual subcomponents in multimodal communication by the frog Allobates femoralis." Anim Behav 79(3): 739-745.
de Rijk, E. P., B. G. Jenks and S. E. Wendelaar Bonga (1990). "Morphology of the pars intermedia and the melanophore-stimulating cells in Xenopus laevis in relation to background adaptation." Gen Comp Endocrinol 79(1): 74-82.
de Rijk, E. P., F. J. van Strien and E. W. Roubos (1992). "Demonstration of coexisting catecholamine (dopamine), amino acid (GABA), and peptide (NPY) involved in inhibition of melanotrope cell activity in Xenopus laevis: a quantitative ultrastructural, freeze-substitution immunocytochemical study." J Neurosci 12(3): 864-871.
DeMartini, E. E. (1985). "Social behavior and coloration changes in painted greenling, Oxylebius pictus (Pisces: Hexagrammidae)." Copeia: 966-975.
Doucet, S. M. and D. J. Mennill (2010). "Dynamic sexual dichromatism in an explosively breeding Neotropical toad." Biol Lett 6(1): 63-66.
Ellingson, J. M., L. J. Fleishman and E. R. Loew (1995). "Visual pigments and spectral sensitivity of the diurnal gecko Gonatodes albogularis." J Comp Physiol A 177(5): 559-567.
Fleishman, L. J., E. R. Loew and M. J. Whiting (2011). "High sensitivity to short wavelengths in a lizard and implications for understanding the evolution of visual systems in lizards." Proc Biol Sci 278(1720): 2891-2899.
Fujii, R. (2000). "The regulation of motile activity in fish chromatophores." Pigment Cell Res 13(5): 300-319.
Gerhardt, H. C., C. C. Martinez-Rivera, J. J. Schwartz, V. T. Marshall and C. G. Murphy (2007). "Preferences based on spectral differences in acoustic signals in four species of treefrogs (Anura: Hylidae)." J Exp Biol 210(Pt 17): 2990-2998.
Gomez, D., C. Richardson, T. Lengagne, M. Derex, S. Plenet, P. Joly, J. P. Lena and M. Thery (2010). "Support for a role of colour vision in mate choice in the nocturnal European treefrog (Hyla arborea)." Behaviour 147(13-14): 1753-1768.
Gomez, D., C. Richardson, T. Lengagne, S. Plenet, P. Joly, J. P. Lena and M. Thery (2009). "The role of nocturnal vision in mate choice: females prefer conspicuous males in the European tree frog (Hyla arborea)." Proc Biol Sci 276(1666): 2351-2358.
Gomez, D., C. Richardson, M. Théry, T. Lengagne, J. P. LÉNA, S. Plénet and P. Joly (2011). "Multimodal signals in male European treefrog (Hyla arborea) and the influence of population isolation on signal expression." Biol J Linn Soc 103(3): 633-647.
Höglund, E., P. H. M. Balm and S. Winberg (2002). "Behavioural and neuroendocrine effects of environmental background colour and social interaction in Arctic charr (Salvelinus alpinus)." J Exp Biol 205(16): 2535-2543.
Hailman, J. P. and R. G. Jaeger (1974). "Phototactic responses to spectrally dominant stimuli and use of colour vision by adult anuran amphibians: a comparative survey." Anim Behav 22: 757-795.
Hamer, R., F. L. Lemckert and P. B. Banks (2011). "Adult frogs are sensitive to the predation risks of olfactory communication." Biol Lett 7(3): 361-363.
Hart, N. S. (2001). "The visual ecology of avian photoreceptors." Prog Retin Eye Res 20(5): 675-703.
Hart, N. S. (2004). "Microspectrophotometry of visual pigments and oil droplets in a marine bird, the wedge-tailed shearwater Puffinus pacificus: topographic variations in photoreceptor spectral characteristics." J Exp Biol 207(7): 1229-1240.
Hart, N. S. and D. M. Hunt (2007). "Avian visual pigments: Characteristics, spectral tuning, and evolution." Am Nat 169(1): S7-S26.
Hart, N. S., J. C. Partridge, A. T. D. Bennett and I. C. Cuthill (2000). "Visual pigments, cone oil droplets and ocular media in four species of estrildid finch." J Comp Physiol A 186(7-8): 681-694.
Hasson, O. (1989). "Amplifiers and the handicap principle in sexual selection: a different emphasis." Proc R Soc Lond B Biol Sci 235(1281): 383-406.
Healey, E. (1999). "The skin pattern of young plaice and its rapid modification in response to graded changes in background tint and pattern." J Fish Biol 55(5): 937-971.
Hettyey, A., G. Herczeg, A. Laurila, P. A. Crochet and J. Merila (2009). "Body temperature, size, nuptial colouration and mating success in male Moor Frogs (Rana arvalis)." Amphibia-Reptilia 30(1): 37-43.
Hoffmann, F. and W. Kloas (2012). "Estrogens Can Disrupt Amphibian Mating Behavior." PLoS One 7(2).
Hu, J. R., J. Z. Du and X. Ji (2004). "Pattern of plasma sex steroid hormone levels during the breeding season of male and female skink: Eumeces chinensis." Shi Yan Sheng Wu Xue Bao 37(6): 443-448.
Ichikawa, Y., H. Ohtani and I. Miura (2001). "The yellow mutation in the frog Rana rugosa: pigment organelle deformities in the three types of chromatophore." Pigment Cell Res 14(4): 283-288.
Ives, J. T., R. A. Normann and P. W. Barber (1983). "Light intensification by cone oil droplets: electromagnetic considerations." JOSA 73(12): 1725-1731.
Jenks, B. G., L. Galas, M. Kuribara, L. Desrues, A. H. Kidane, H. Vaudry, W. J. Scheenen, E. W. Roubos and M. C. Tonon (2011). "Analysis of the melanotrope cell neuroendocrine interface in two amphibian species, Rana ridibunda and Xenopus laevis: a celebration of 35 years of collaborative research." Gen Comp Endocrinol 170(1): 57-67.
Jenks, B. G., A. H. Kidane, W. J. Scheenen and E. W. Roubos (2007). "Plasticity in the melanotrope neuroendocrine interface of Xenopus laevis." Neuroendocrinology 85(3): 177-185.
Jessop, T. S., R. Chan and D. Stuart-Fox (2009). "Sex steroid correlates of female-specific colouration, behaviour and reproductive state in Lake Eyre dragon lizards, Ctenophorus maculosus." J Comp Physiol A Neuroethol Sens Neural Behav Physiol 195(7): 619-630.
Johnson, M. L., E. Gaten and P. M. J. Shelton (2002). "Spectral sensitivities of five marine decapod crustaceans and a review of spectral sensitivity variation in relation to habitat." J Mar Biol Assoc UK 82(5): 835-842.
Kelber, A., A. Balkenius and E. J. Warrant (2002). "Scotopic colour vision in nocturnal hawkmoths." Nature 419(6910): 922-925.
Kelber, A. and L. S. V. Roth (2006). "Nocturnal colour vision–not as rare as we might think." J Exp Biol 209(5): 781-788.
Kempenaers, B. (2007). "Mate choice and genetic quality: a review of the heterozygosity theory." Adv Study Behav 37: 189-278.
Kidane, A. H., P. M. Cruijsen, M. A. Ortiz-Bazan, H. Vaudry, J. Leprince, F. J. Kuijpers-Kwant, E. W. Roubos and B. G. Jenks (2007a). "Actions of PACAP and VIP on melanotrope cells of Xenopus laevis." Peptides 28(9): 1790-1796.
Kidane, A. H., G. Heinrich, R. P. Dirks, B. A. de Ruyck, N. H. Lubsen, E. W. Roubos and B. G. Jenks (2009). "Differential neuroendocrine expression of multiple brain-derived neurotrophic factor transcripts." Endocrinol 150(3): 1361-1368.
Kidane, A. H., E. W. Roubos and B. G. Jenks (2008). "Pituitary adenylate cyclase-activating polypeptide regulates brain-derived neurotrophic factor exon IV expression through the VPAC1 receptor in the amphibian melanotrope cell." Endocrinol 149(8): 4177-4182.
Kidane, A. H., S. H. van Dooren, E. W. Roubos and B. G. Jenks (2007b). "Expression and physiological regulation of BDNF receptors in the neuroendocrine melanotrope cell of Xenopus laevis." Gen Comp Endocrinol 153(1-3): 176-181.
King, R. B., J. K. Douglass, J. B. Phillips and C. L. Baube (1993). "Scotopic spectral sensitivity of the optomotor response in the green treefrog Hyla cinerea." J Exp Zool 267(1): 40-46.
Kirschfeld, K. (1982). "Carotenoid pigments: their possible role in protecting against photooxidation in eyes and photoreceptor cells." Proc R Soc Lond B Biol Sci 216(1202): 71-85.
Kodric-Brown, A. (1998). "Sexual dichromatism and temporary color changes in the reproduction of fishes." Amer Zool 38(1): 70-81.
Kokko, H., R. Brooks, J. M. McNamara and A. I. Houston (2002). "The sexual selection continuum." Proc Biol Sci 269(1498): 1331-1340.
Kram, Y. A., S. Mantey and J. C. Corbo (2010). "Avian cone photoreceptors tile the retina as five independent, self-organizing mosaics." PLoS One 5(2): e8992.
Kramer, B. M. R., P. M. J. M. Cruijsen, D. T. W. M. Ouwens, M. W. Coolen, G. J. M. Martens, E. W. Roubos and B. G. Jenks (2002). "Evidence that brain-derived neurotrophic factor acts as an autocrine factor on pituitary melanotrope cells of Xenopus laevis." Endocrinol 143(4): 1337-1345.
Kuribara, M., M. W. Hess, M. Cazorla, E. W. Roubos, W. J. Scheenen and B. G. Jenks (2011). "Brain-derived neurotrophic factor stimulates growth of pituitary melanotrope cells in an autocrine way." Gen Comp Endocrinol 170(1): 156-161.
Kuribara, M., A. H. Kidane, G. A. Vos, D. de Gouw, E. W. Roubos, W. J. Scheenen and B. G. Jenks (2011). "Extracellular-signal regulated kinase regulates production of pro-opiomelanocortin in pituitary melanotroph cells." J Neuroendocrinol 23(3): 261-268.
Larsson, D. G., K. Kinnberg, J. Sturve, E. Stephensen, M. Skon and L. Forlin (2002). "Studies of masculinization, detoxification, and oxidative stress responses in guppies (Poecilia reticulata) exposed to effluent from a pulp mill." Ecotoxicol Environ Saf 52(1): 13-20.
Larsson, D. G. J., K. Kinnberg, J. Sturve, E. Stephensen, M. Skon and L. Forlin (2002). "Studies of masculinization, detoxification, and oxidative stress responses in guppies (Poecilia reticulata) exposed to effluent from a pulp mill." Ecotoxicol Environ Saf 52(1): 13-20.
Leary, C. J., D. J. Fox, D. B. Shepard and A. M. Garcia (2005). "Body size, age, growth and alternative mating tactics in toads: satellite males are smaller but not younger than calling males." Anim Behav 70: 663-671.
Leary, C. J., T. S. Jessop, A. M. Garcia and R. Knapp (2004). "Steroid hormone profiles and relative body condition of calling and satellite toads: implications for proximate regulation of behavior in anurans." Behav Ecol 15(2): 313-320.
Leclaire, S., J. White, E. Arnoux, B. Faivre, N. Vetter, S. A. Hatch and E. Danchin (2011). "Integument coloration signals reproductive success, heterozygosity, and antioxidant levels in chick-rearing black-legged kittiwakes." Naturwissenschaften 98(9): 773-782.
Leclercq, E., J. F. Taylor and H. Migaud (2010). "Morphological skin colour changes in teleosts." Fish and Fisheries 11(2): 159-193.
Lima, M. S. C. S. and J. Pederassi (2012). "Morphometrics and ratio of body proportionality of tadpoles of Rhinella icterica (Anura, Bufonidae) at different developmental stages." Brazil J Biol 72(3): 623-629.
Loew, E. R., L. J. Fleishman, R. G. Foster and I. Provencio (2002). "Visual pigments and oil droplets in diurnal lizards: a comparative study of Caribbean anoles." J Exp Biol 205(Pt 7): 927-938.
Loew, E. R. and V. I. Govardovskii (2001). "Photoreceptors and visual pigments in the red-eared turtle, Trachemys scripta elegans." Vis Neurosci 18(5): 753-757.
Loew, E. R., V. I. Govardovskii, P. Rohlich and A. Szel (1996). "Microspectrophotometric and immunocytochemical identification of ultraviolet photoreceptors in geckos." Vis Neurosci 13(2): 247-256.
Lopez-Lopez, R., M. Lopez-Gallardo, M. J. Perez-Alvarez and C. Prada (2008). "Isolation of chick retina cones and study of their diversity based on oil droplet colour and nucleus position." Cell Tissue Res 332(1): 13-24.
Lopez, R., M. Lopez-Gallardo, I. Busturia, L. Anezary and C. Prada (2005). "Spatial and temporal patterns of growth and differentiation of cone oil droplets in the chick retina." J Neurosci Res 79(3): 401-411.
Lozano, G. A. (1994). "Carotenoids, parasites, and sexual selection." Oikos: 309-311.
Lu, X., W. Chen, L. X. Zhang and X. Y. Ma (2010). "Mechanical constraint on size-assortative paring success in a temperate frog: An experimental approach." Behav Processes 85(2): 181-184.
Lu, X., X. Y. Ma, L. Q. Fan and T. L. Yu (2010). "Opportunity for sexual selection arises at moderate densities in the frog Rana chensinensis: an experimental approach." J Ethol 28(2): 257-262.
Macedonia, J. M., J. F. Husak, Y. M. Brandt, A. K. Lappin and T. Baird (2004). "Sexual dichromatism and color conspicuousness in three populations of collared lizards (Crotaphytus collaris) from Oklahoma." J of Herpetol38(3): 340-354.
Marquez, R., J. Bosch and X. Eekhout (2008). "Intensity of female preference quantified through playback setpoints: call frequency versus call rate in midwife toads." Anim Behav 75: 159-166.
Martin, J. and P. Lopez (2010). "Multimodal sexual signals in male ocellated lizards Lacerta lepida: vitamin E in scent and green coloration may signal male quality in different sensory channels." Naturwissen 97(6): 545-553.
Martin, T. E. and A. V. Badyaev (1996). "Sexual dichromatism in birds: Importance of nest predation and nest location for females versus males." Evol 50(6): 2454-2460.
Matthews, H. R. and A. P. Sampath (2010). "Photopigment quenching is Ca2+ dependent and controls response duration in salamander L-cone photoreceptors." J Gen Physiol 135(4): 355-366.
McCord, C. P. and F. P. Allen (2005). "Evidences associating pineal gland function with alterations in pigmentation." J Exp Zool 23(1): 207-224.
Miller, G. F. (1999). "Sexual selection for cultural displays." Evol cul: 71-91.
Moller, A. and A. Pomiankowski (1993). "Why have birds got multiple sexual ornaments?" Behav Ecol Sociobiol 32(3): 167-176.
Moore, M. C. (1986). "Elevated testosterone levels during nonbreeding-season territoriality in a fall-breeding lizard, Sceloporus jarrovi." J Comp Physiol A 158(2): 159-163.
Moravec, J., J. Aparicio, M. Guerrero-Reinhard, G. Calderon, K. H. Jungfer and V. Gvozdik (2009). "A new species of Osteocephalus (Anura: Hylidae) from Amazonian Bolivia: first evidence of tree frog breeding in fruit capsules of the Brazil nut tree." Zootaxa(2215): 37-54.
Moravec, J., J. Aparicio and J. Kohler (2006). "A new species of tree frog, genus Dendropsophus (Anura : Hylidae), from the Amazon of northern Bolivia." Zootaxa(1327): 23-40.
Murphy, C. G. (1994). "Chorus tenure of male barking treefrogs, Hyla gratiosa." Anim Behav 48(4): 763-777..
Noriega, N. C. and T. B. Hayes (2000). "DDT congener effects on secondary sex coloration in the reed frog Hyperolius argus: a partial evaluation of the Hyperolius argus endocrine screen." Comp Biochem Physiol B Biochem Mol Biol 126(2): 231-237.
Olson, V. A. and I. P. Owens (1998). "Costly sexual signals: are carotenoids rare, risky or required?" Trends Ecol Evol 13(12): 510-514.
Oshima, N., M. Makino, S. Iwamuro and H. A. Bern (1996). "Pigment dispersion by prolactin in cultured xanthophores and erythrophores of some fish species." J Exp Zool 275(1): 45-52.
Osorno, J. L., A. Nunez-de la-Mora, L. D'Alba and J. C. Wingfield (2010). "Hormonal correlates of breeding behavior and pouch color in the Magnificent Frigatebird, Fregata magnificens." Gen Comp Endocrinol 169(1): 18-22.
Pellegri, G., P. J. Magistretti and J. L. Martin (1998). "VIP and PACAP potentiate the action of glutamate on BDNF expression in mouse cortical neurones." Eur J Neurosci 10(1): 272-280.
Pryke, S. R., S. Andersson and M. J. Lawes (2001). "Sexual selection of multiple handicaps in the red-collared widowbird: female choice of tail length but not carotenoid display." Evol 55(7): 1452-1463.
Raffin-Sanson, M. L., Y. de Keyzer and X. Bertagna (2003). "Proopiomelanocortin, a polypeptide precursor with multiple functions: from physiology to pathological conditions." Eur J Endocrinol 149(2): 79-90.
Richardson, C., J. P. Lena, P. Joly and T. Lengagne (2008). "Are leaders good mates? A study of call timing and male quality in a chorus situation." Anim Behav 76: 1487-1495.
Richardson, C., J. Popovici, F. Bellvert and T. Lengagne (2009). "Conspicuous colouration of the vocal sac of a nocturnal chorusing treefrog: carotenoid-based?" Amphibia-Reptilia 30(4): 576-580.
Rijli, F. M., S. De Lucchini, G. Ciliberto and G. Barsacchi (1993). "A Zn-finger protein, Xfin, is expressed during cone differentiation in the retina of the frog Xenopus laevis." Int J Dev Biol 37(2): 311-317.
Roth, L. S. and A. Kelber (2004). "Nocturnal colour vision in geckos." Proc Biol Sci 271 Suppl 6: S485-487.
Roubos, E. W., W. J. Scheenen and B. G. Jenks (2005). "Neuronal, neurohormonal, and autocrine control of Xenopus melanotrope cell activity." Ann N Y Acad Sci 1040: 172-183.
Ryan, M. J., X. E. Bernal and A. S. Rand (2010). "Female mate choice and the potential for ornament evolution in tungara frogs Physalaemus pustulosus." Cur Zool 56(3): 343-357.
Ryan, M. J. and A. Keddy-Hector (1992). "Directional patterns of female mate choice and the role of sensory biases." American Naturalist: 4-35.
Ryser, J. (1989). "Weight loss, reproductive output, and the cost of reproduction in the common frog, Rana temporaria." Oecologia 78(2): 264-268.
Sage, M. (1970). "Control of prolactin release and its role in color change in the teleost Gillichthys mirabilis." J Exp Zool 173(1): 121-127.
Sherman, C. D. H., J. Sagvik and M. Olsson (2010). "Female Choice for Males with Greater Fertilization Success in the Swedish Moor Frog, Rana arvalis." PLoS One 5(10).
Skold, H. N., T. Amundsen, P. A. Svensson, I. Mayer, J. Bjelvenmark and E. Forsgren (2008). "Hormonal regulation of female nuptial coloration in a fish." Horm Behav 54(4): 549-556.
Steffen, J. E. (2007). Carotenoid-based Dewlap Color as a Visual Signal in Social Communication of Brown Anoles (Norops Sagrei), ProQuest.
Stelkens, R. B., M. E. Pierotti, D. A. Joyce, A. M. Smith, I. van der Sluijs and O. Seehausen (2008). "Disruptive sexual selection on male nuptial coloration in an experimental hybrid population of cichlid fish." Philos Trans R Soc Lond B Biol Sci 363(1505): 2861-2870.
Sugden, D., K. Davidson, K. A. Hough and M. T. Teh (2004). "Melatonin, melatonin receptors and melanophores: a moving story." Pigment Cell Res 17(5): 454-460.
Summers, K., R. Symula, M. Clough and T. Cronin (1999). "Visual mate choice in poison frogs." Proc Biol Sci 266(1434): 2141-2145.
Svensson, P. A., E. Forsgren, T. Amundsen and H. N. Skold (2005). "Chromatic interaction between egg pigmentation and skin chromatophores in the nuptial coloration of female two-spotted gobies." J Exp Biol 208(Pt 23): 4391-4397.
Sztatecsny, M., D. Preininger, A. Freudmann, M. C. Loretto, F. Maier and W. Hodl (2012). "Don't get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations." Behav Ecol Sociobiol 66(12): 1587-1593.
Sztatecsny, M., C. Strondl, A. Baierl, C. Ries and W. Hodl (2010). "Chin up: are the bright throats of male common frogs a condition-independent visual cue?" Anim Behav 79(4): 779-786.
Takahashi, D. (2000). "Conventional sex roles in an amphidromous Rhinogobius goby in which females exhibit nuptial coloration." Ichthyol Res 47(3): 303-306.
Todd, B. D. and A. K. Davis (2007). "Sexual dichromatism in the marbled salamander Ambyastoma opacum." Can J Zool 85(9): 1008-1013.
Toft, G. and E. Baatrup (2001). "Sexual characteristics are altered by 4-tert-octylphenol and 17beta-estradiol in the adult male guppy (Poecilia reticulata)." Ecotoxicol Environ Saf 48(1): 76-84.
Toft, G. and E. Baatrup (2003). "Altered sexual characteristics in guppies (Poecilia reticulata) exposed to 17 beta-estradiol and 4-tert-octylphenol during sexual development." Ecotoxicol Environ Saf 56(2): 228-237.
Toyoda, F., I. Hasunuma, K. Yamamoto, M. Yamashita and S. Kikuyama (2005). "Prolactin acts centrally to enhance newt courtship behavior." Gen Comp Endocrinol 141(2): 172-177.
Tsuboi, M., A. Gonzalez-Voyer, J. Hoglund and N. Kolm (2012). "Ecology and mating competition influence sexual dimorphism in Tanganyikan cichlids." Evol Ecol 26(1): 171-185.
Ubink, R., R. Tuinhof and E. W. Roubos (1998). "Identification of suprachiasmatic melanotrope-inhibiting neurons in Xenopus laevis: a confocal laser-scanning microscopy study." J Comp Neurol 397(1): 60-68.
Vasquez, T. and K. S. Pfennig (2007). "Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, Scaphiopus couchii." Behav Ecol Sociobiol 62(1): 127-135.
Vaudry, D., A. Falluel-Morel, S. Bourgault, M. Basille, D. Burel, O. Wurtz, A. Fournier, B. K. Chow, H. Hashimoto, L. Galas and H. Vaudry (2009). "Pituitary adenylate cyclase-activating polypeptide and its receptors: 20 years after the discovery." Pharmacol Rev 61(3): 283-357.
Vazquez-Martinez, R., J. R. Peinado, J. L. Gonzalez De Aguilar, L. Desrues, M. C. Tonon, H. Vaudry, F. Gracia-Navarro and M. M. Malagon (2001). "Melanotrope cell plasticity: a key mechanism for the physiological adaptation to background color changes." Endocrinol 142(7): 3060-3067.
von Schantz, T., S. Bensch, M. Grahn, D. Hasselquist and H. Wittzell (1999). "Good genes, oxidative stress and condition-dependent sexual signals." Proc Biol Sci 266(1414): 1-12.
Vorobyev, M. (2003). "Coloured oil droplets enhance colour discrimination." Proc Biol Sci 270(1521): 1255-1261.
Vorobyev, M. (2004). "Ecology and evolution of primate colour vision." Clin Exp Optom 87(4-5): 230-238.
Walls, G. L. and H. Judd (1933). "The intra-ocular colour-filters of vertebrates." Br J Ophthalmol 17(11): 641.
Walsh, N., J. Dale, K. J. McGraw, M. A. Pointer and N. I. Mundy (2012). "Candidate genes for carotenoid coloration in vertebrates and their expression profiles in the carotenoid-containing plumage and bill of a wild bird." Proc Biol Sci 279(1726): 58-66.
Wells, K. D. and T. L. Taigen (1989). "Calling Energetics of a Neotropical Treefrog, Hyla microcephala." Behav Ecol Sociobiol: 13-22.
Wogel, H., P. A. Abrunhosa and J. P. Pombal (2005). "Breeding behaviour and mating success of Phyllomedusa rohdei (Anura, Hylidae) in south-eastern Brazil." J Nat Hist 39(22): 2035-2045.
Woodgate, J. L., A. T. D. Bennett, S. Leitner, C. K. Catchpole and K. L. Buchanan (2010). "Developmental stress and female mate choice behaviour in the zebra finch." Anim Behav 79(6): 1381-1390.
Yokoyama, S. (2000). "Molecular evolution of vertebrate visual pigments." Prog Retin Eye Res 19(4): 385-419.
Young, S. and G. Martin (1984). "Optics of retinal oil droplets: a model of light collection and polarization detection in the avian retina." Vision Res 24(2): 129-137.
Zimmermann, M. (1983). "Ethical guidelines for investigations of experimental pain in conscious animals." Pain 16(2): 109.
林中一 (2001). "台北縣雙溪河流域褐樹蛙 (Buergeria robusta) 之生殖生態與族群分布." 國立台灣大學動物研究所碩士論文 48頁.
楊懿如 (2006). "太魯閣國家公園兩棲類調查及監測計畫報告."