University of Groningen. Maternal phenotypic engineering Müller, Wendt

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1 University of Groningen Maternal phenotypic engineering Müller, Wendt IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2004 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Müller, W. (2004). Maternal phenotypic engineering: Adaptation and constraint in prenatal maternal effects Groningen: s.n. Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date:

2 References References 171

3 Banks, E.M Social organization in red jungle fowl hens (Gallus gallus subsp.). Ecology 37: Barua, A., Yoshimura, Y & Tamura, T Effects of ageing and oestrogens on the localization of immunoglobulin-containing cells in the chicken ovary. J. Reprod. Fertil. 114: Batty, K.A., Herbert, J., Keverne, E.B. & S.V. Vellucci, Differences in blood levels of androgens in female tatapoin monkeys related to their social status. Neuroendocrinology 44: Behnke, J.M., Barnard, C.J. & D. Wakelin, Understanding chronic nematode infections: evolutionary considerations current hypotheses and the way forward. Int. J. Parasitol. 22: Beletsky, L.D., Orians, G.H. & J.C. Wingfield, Year-to-year patterns of circulating levels of testosterone and corticosterone in relation to breeding density, experience, and reproductive success of the polygynous Red-winged Blackbird. Horm. Behav. 26: Beukeboom, L., Dijkstra, C., Daan, S. & T. Meijer, Seasonality of clutch size determination in the kestrel, Falco tinnunculus: an experimental study. Ornis Scand. 19: Blank, J.L. & V. Nolan, Offspring sex ratio in red-winged blackbirds is dependent on maternal age. Proc. Natl. Acad. Sci. USA 80: Blount, J.D., Møller, A.P. & D.C. Houston, Why egg yolk is yellow. TREE 15: Blount, J.D., Surai, P.F., Nager, R.G., Houston, D.C., Møller, A.P., Trewby, M.L. & M. Kennedy, Carotenoids and egg quality in the lesser black-backed gull Larus fuscus: a supplemental feeding study of maternal effects. Proc. R. Soc. Lond B 269: Blount, J.D., Metcalfe, N.B., Birkhead, T.R. & P.F. Surai, Carotenoid modulation of immune function and sexual attractiveness in zebra finches. Science 300: Blount, J.D., Houston, D.C., Surai, P.F. & A.P. Møller, Egg laying capacity is limited by carotenoid pigment availability in wild gulls Larus fuscus. Proc. R. Soc. Lond B, 271 (Suppl. 3): Bollinger, P.B., Bollinger, E.K., & R.A. Malecki, Tests of the three hypotheses of hatching asynchrony in the common tern. Auk 107: Bolton, M Determinants of chick survival in the lesser black-backed gull: relative contributions of egg size and parental quality. J. Anim. Ecol. 61: Bolton, M., Houston, D. & P. Monaghan, Nutritional constraints on egg formation in the lesser black-backed gull: An experimental study. J. Anim. Ecol. 61: Bortolotti, G.R Influence of sibling competition on nestling sex ratio in sexually dimorphic birds. Am. Nat. 127: References 173

4 Bowden, R.M., Ewert, M.A. & C.E. Nelson, Environmental sex determination in a reptile varies seasonally and with yolk hormones. Proc. R. Soc. Lond. B, 267: Bowry, T.R Immunology. Oxford University Press, Oxford. Bradbury, R.R. & J.K. Blakey, Diet, maternal condition, and offspring sex ratio in the zebra finch, Poephila guttata. Proc. R. Soc. Lond. B 265: Bradbury, R.B. & R. Griffiths, Sex-biased mortality is influenced by hatching asynchrony in the Lesser Black-backed Gull Larus fuscus. J. Avian Biol. 30: Braude, S., Tang-Martinez, Z. & G.T. Taylor, Stress, testosterone, and the immunoredistribution hypothesis. Behav. Ecol. 10: Brommer, J.E Immunocompetence and its costs during development: an experimental study in blue tit nestlings. Proc. R. Soc. Lond. B 271 (Suppl. 3): Brouwer, A. & A.L. Spaans, Egg predation in the Herring Gull Larus argentatus: why does it vary so much between nests? Ardea 82: Brown C.R. & M.B.Brown, Ectoparasitism as a cost of coloniality in cliff swallows (Hirundo pyrrhonota). Ecology 67: Brush, A.H Metabolism of carotenoid pigments in birds. FASEB Journal 4: Buechler, K., Fitze, P.S., Gottstein, B., Jacott, A. & H. Richner, Parasiteinduced maternal response in a natural bird population. J. Anim Ecol. 71: Buntin, J.D., Advis, J.P., Ottinger, M.A., Lea, R.W. & P.J. Sharp, An analysis of physiological mechanisms underlying the antigonadotropic action of intracranial prolactin in ringdoves. Gen. Comp. Endocrinol. 114: Burke, W.H Effects of in ovo injection of an antiandrogen on embryonic and posthatching growth of broiler chicks. Poultry Science, 75 : C Casto, J.M., Nolan, V. & E.D. Ketterson, Steroid hormones and immune function: Experimental studies in wild and captive dark-eyed juncos (Junco hyemalis). Am. Nat. 157: Charnov, E.L The genetical evolution of patterns of sexuality: Darwinian fitness. Am. Nat. 113: Charnov, E.L The theory of sex allocation. Princeton, Princeton University press. Christe, P., Møller, A.P. & F. de Lope, Immunocompetence and nestling survival in the house martin: the tasty chick hypothesis. Oikos 83: Christians, J.K Avian egg size: variation within species and inflexibility within individuals. Biol. Rev. 77:

5 Clark, A.B. & D.S. Wilson, The onset of incubation in birds. Am Nat 125: Clark, A.B. & D.S. Wilson, Avian breeding adaptations: hatching asynchrony, brood reduction and nest failure. Q. Rev. Biol. 56: Clark, M.M. & B.G. Jr. Galef., Prenatal influences on reproductive life history strategies. TREE 10: Clifton, P.G. & R.J. Andrew, Contrasting effects of pre-hatch and posthatch exposure to gonadal-steroids on the development of vocal, sexual, and aggressive behavior of young domestic fowl. Horm. Behav. 23: Clinton M. & Haines L.C An overview of factors influencing sex determination and gonadal development in birds. Cell Mol. Life Sci. 55: Clutton-Brock, T. H., Albon, S. D. & F. E. Guinness, Maternal dominance, breeding success and birth sex ratios in red deer. Nature 308: Clutton-Brock, T.H., Albon S.D. & F.E. Guinness, Parental investment and sex differences in juvenile mortality in birds and mammals. Nature 313: Clutton-Brock, T.H The evolution of parental care. Princeton university press, Princeton, NJ. Collias, N.E Statistical analysis of factors which make for success in initial encounters between hens. Am. Nat. 77: Collias, N.E., Collias, E.C. & R.I. Jennrich, Dominant red junglefowl hens in an unconfined flock rear the most young over their lifetime. Auk 111: Cordero, P.J., Griffith, S.C., Aparicio, J.M. & D.T. Parkin, Sexual dimorphism in house sparrow eggs. Behav. Ecol. Sociobiol. 48: Cordero, P.J., Viñuela, J., Aparicio, J.M. & J.P. Veiga, Seasonal variation in sex ratio and sexual egg dimorphism favouring daughters in first clutches of the spotless starling. J. Evol. Biol. 14: Coulson, P.B., Thornthwaite, J.T., Skafar, D.F. & S.S. Seaver, Modulation of glucocorticoid hormone receptor levels in chicken lymphoid tissue following treatment with androgens in vivo. J. Steroid Biochem. 17: 1-9. Courtney, P., Seasonal variation among intra-clutch hatching intervals among common tern Sterna hirundo. Ibis 121: Cramp, S. & K.E.L. Simmons, (eds.) Handbook of the birds of Europe, the Middle East and North Africa: the birds of the Western Paleartic. Volume III: waders to gulls. Oxford University Press, Oxford. Creel, S Social dominance and stress hormones. TREE 16: D Danchin, E. & R.H. Wagner, The evolution of coloniality: the emergence of new perspectives. TREE 12: References 175

6 David, P., Bjorksten, T., Fowler, K. & A. Pomiankowski, Conditiondependent signalling of genetic variation in stalk-eyed flies. Nature 406: De Kogel, C.H. & H.J. Prijs, Effects of brood size manipulations on sexual attractiveness of offspring in the zebra finch. Anim. Behav. 51: Dijkstra, C., Daan, S. & J.B. Buker, Adaptive seasonal variation in the sex ratio of kestrel broods. Funct. Ecol. 4: Dijkstra, C., Daan, S. & I. Pen, Fledging sex ratios in relation to brood size in size-dimorphic altricial birds. Behav. Ecol. 9: Dohms, J.E., Saif, Y.M. & W.L. Bacon Metabolism and passive transfer of immunoglobulins in the turkey hen. A. J. Vet. Res. 39: Drummond, H., Osorno, J.L., Torres, R., Garcia Chavelas, C. & H. Merchant Larios, Sexual size dimorphism and sibling competition: implications for avian sex ratios. Am. Nat. 138: Duffy, D.L., Bentley, G.E., Drazen, D.L. & G.F. Ball, Effects of testosterone on cell-mediated and humoral immunity in non-breeding adult European starlings. Behav. Ecol. 11: Dunlop, E On incubation. British Birds 4: Dzus, E.H., Bortolotti, G.R. & J.M. Gerrard, Does sex-biased hatching order in bald eagles vary with food resources? Ecoscience 3: E Eising, C.M., Eikenaar, C., Schwabl, H. & T.G.G. Groothuis, Maternal androgens in black-headed gull (Larus ridibundus) eggs: consequences for chick development. Proc. R. Soc. Lond B 268: Eising, C.M. & T.G.G. Groothuis, Yolk androgens and begging behavior in black-headed gull chicks: and experimental field study. Anim. Behav. 66: Eising, C.M., Visser, G.H., Müller, W. & T.G.G. Groothuis, Steroids for free? No metabolic costs of elevated maternal androgen levels in the blackheaded gull. J. Exp. Biol. 206: Eising, C., Müller, W., Dijkstra, C. & T.G.G. Groothuis, Maternal androgens in egg yolks: relation with sex, incubation time and embryonic growth. Gen. Comp. Endocrinol. 132: Eising, C.M Mothers knows best? Costs and benefits of differential maternal hormone allocation in birds. PhD thesis University of Groningen. Elf, P.K. & A.J. Fivizanni, Changes in sex steroid levels in yolks of the Leghorn chicken, Gallus domesticus, during embryonic development. J. Exp. Zool. 293: Ellis, L.A., Borst, D.W. & C.F. Thompson, Hatching asynchrony and maternal androgens in egg yolks of house wrens. J. Avian Biol. 32:

7 Evans, R.M Embryonic fine tuning of pipped egg temperature in the American white pelican. Anim. Behav. 40: F Fargallo, A.J., Laaksonen, T., Pöyri, V. & E. Korpimäki, Inter-sexual differences in the immune response of Eurasian kestrel nestlings under food shortage. Ecology Letters 5: Fasler, R., Schwartz, S., Dietrich, H. & G. Wick, Sex steroid and glucocorticoid receptors in the bursa of Fabricius of obese strain chickens spontaneously developing autoimmune thyroiditis. J. Steroid Biochem. 24: Folstad, I. & A.J. Carter, Parasites, bright males, and the immunocompetence handicap. Am. Nat. 139: Forbes, L.S The good, the bad and the ugly: Lack s brood reduction hypothesis and experimental design. J. Avian Biol. 25: Forbes, L.S., Thornton, S., Glassey, B., Forbes, M. & N. Buckley, Why parent birds play favourites. Nature 390: Frank, L.G., Davidson, J.M. & E.R. Smith, Androgen levels in the spotted hyena Crocuta crocuta: the influence of social factors. J. Zool. Lond. A 206: French, J.B. Jr., Nisbet, I.C.T. & H. Schwabl, Maternal steroids and contaminants in common tern eggs: a mechanism of endocrine disruption? Comp. Biochem. Physiol. 128: G Gabrielsen, G.W., Klaassen, M. & F. Mehlum, Energetics of black-legged kittiwake Rissa Tridactyla chicks. Ardea 80: Gasparini, J., McCoy K.D., Tveraa T. & T. Boulinier, Induced maternal response to the Lyme disease spirochaete Borrelia burgdorferi sensu lato in a colonial seabird the kittiwake Rissa tridactyla. Proc. R. Soc. Lond. B 268: Gebhardt-Henrich, S. & H. Richner, Causes of growth variation and its consequences for fitness. In: Starck, J.M., Ricklefs, R.E. (eds): Avian growth and development evolution within the altricial-precocial. Oxford University Press, Oxford, Chapter 8, pp Gibbons, D.W Hatching asynchrony reduces parental investment in the jackdaw. J. Anim. Ecol. 56: Gil, D., Graves, J., Hazon, N. & A. Wells, Male attractiveness and differential testosterone investment in zebra finch eggs. Science 286: Glick, B., Bursa of Fabricius. In: Avian biology Vol.7, pp , eds Farner DS, King JR Parkes KC, Academic Press, New York. References 177

8 178 Glutz von Blotzheim, U.N. & K.M. Bauer, Handbuch der Vögel Mitteleuropas. Band 8. Wiesbaden. Götemark, F. & M. Andersson, Colonial breeding reduces nest predation in the common gull (Larus canus). Anim. Behav. 32: Godfray, H.C.J Evolutionary theory of parent-offspring conflict. Nature 376: Godsave, S.F., Lohmann, R., Vloet, R.P.M. & M. Gahr, Androgen receptors in the embryonic zebra finch hindbrain suggest a function for maternal androgens in the perihatching survival. J. Comp. Neurol. 453: Goldstein, H Multilevel statistical approach. Edward Arnold, London. Goldsmith, A.R Prolactin in avian reproductive cycles. In Balthazart J, Pröve E, Gilles R (eds) Hormones and behaviour in higher vertebrates, pp Springer Verlag, Berlin. Grahn, M. & T. von Schantz, Fashion and age in pheasants: age differences in mate choice. Proc. R. Soc. Lond B 255: Granado, F., Olmedilla, B., Gil-Martinez, E. & I. Blanco, Lutein ester in serum after supplementation in human subjects. Br. J. Nutr. 80: Graves, H.B., Hable, C.P. & T.H. Jenkins, Effects of morphology and dominance on female spatial behavior. Behav. Proc. 11: Graves, J., Whiten, A. & P. Henzi, Why does the herring gull lay three eggs? Anim. Behav. 32: Green, D.J., Pair bond influences paternal provisioning and the primary sex ratio of brown thornbills. Anim. Behav. 64: Griffiths, R Sex-biased mortality in the lesser black-backed gull Larus fuscus during the nestling stage. Ibis, 134: Griffiths, R., Double, M.C., Orr K. & R.J.G. Dawson, A DNA test to sex most birds. Mol. Ecol. 7: Grindstaff, J.L., Brodie III, E.D. & E.D. Ketterson, Immune function across generations: integrating mechanism and evolutionary process in maternal antibody transmission. Proc. R. Soc. Lond. B 270: Groothuis, T.G.G. & G. Meeuwissen, The influence of testosterone on the development and fixation of the form of displays in two age classes of young black-headed gulls. Anim. Behav. 43: Groothuis, T.G.G. & H. Schwabl, The influence of laying sequence and habitat characteristics on maternal yolk hormone levels. Funct. Ecol. 16: Grossman, C.J., Interactions between the gonadal steroids and the immune system. Science, 227: Guhl, A.M. & D.C. Warren, Number of offspring sired by cockerels related to social dominance in chickens. Poultry Sci. 25: Guhl, A.M The behaviour of chickens, In: The behaviour of domestic animals (ed. Haffrez). Balliere, Tindall and Cox, London.

9 Gustafsson, L., Quarnström, A. & B. Sheldon, Trade-offs between lifehistory traits and a secondary sexual character in male collared flycatchers. Nature 375: H Hario, M. & E. Rudbäck, Effects of supplementary feeding on the C-egg constituents in the nominate lesser black-backed gull Larus f. fuscus. Ornis Fennica 74: Hario, M. & E. Rudbäck, Dying in the midst of plenty the third chick fate in nominate lesser black-backed gulls Larus f. fuscus. Ornis Fennica 76: Harlow, E. & Lane, D Using antibodies: A laboratory manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY. Hasselquist, D., Marsh, J.A., Sherman, P.W. & J.C. Wingfield, Is avian humoral immunocompetence suppressed by testosterone? Behav. Ecol. Sociobiol. 45: Haq, A., Bailey, C.A. & A. Chinnah, Effect of β-carotene, canthaxanthin, lutein and Vitamin E on neonatal immunity of chicks when supplemented in broiler diets. Poultry Sci. 75: Hebert, P.N. & R. McNeil, Hatching asynchrony and food stress in Ringbilled Gulls: an experimental study. Can. J. Zool. 77: Hegner, R.E. & J.C.Wingfield, Behavioral and endocrine correlates of multiple brooding in the semi-colonial house sparrow. Part II: females. Horm. Behav. 20: Heller E.D., Leitner, G., Drabkin N. & D. Melamed, Passive immunization of chicks against Escherichia coli. Avian Pathol. 19: Henry, M.H. & W.H. Burke, Sexual dimorphism in broiler chick embryos and embryonic muscle development in late incubation. Poultry Sci. 77: Henry, M.H. & W.H. Burke, The effects of in ovo administration of testosterone or an antiandrogen on growth of chick embryos and embryonic muscle characteristics. Poultry Sci. 78: Hiom, L., Bolton, M., Monaghan, P. & D. Worrall, Experimental evidence for food limitation of egg production in gulls. Ornis Scand. 22: Hirota, Y., Suzuki, T., Chazono, Y. & Y. Bito, Humoral immune response characteristics of testosterone-propionate-treated chickens. Immunology 30: Hõrak, P., Tegelmann, L., Ots, I. & A.P. Møller, Immune function and survival of great tit nestlings in relation to growth conditions. Oecologia 121: Hudson, L. & F.C. Hay, Practical immunology. Blackwell Scientific, Oxford, UK. References 179

10 I Iacovides, S. & R.M. Evans, Begging as graded signals of need for food in young ring-billed gulls. Anim. Behav. 56, J Janeway, C.A. & P. Travers, Immunobiology: the immune system in health and disease. 4 th ed. Current biology, London. Jennings, A.R. & E.J.L. Soulsby, Disease in a colony of Black-headed Gulls Larus ridibundus. Ibis 100: Jennings, D.H., Weiss, S.L. & M.L. Moore, Ontogenetic changes in embryonic yolk steroid content in tree lizards: transfer of hormones from the developing embryo to the yolk? Am. Zool. 40: K Kalmbach, E., Nager, R.G., Griffiths, R. & R.W. Furness, Increased reproductive effort results in male-biased offspring sex ratios: an experimental study in a species with reversed sexual size dimorphism. Proc. R. Soc. Lond. B 268: Kariyawasam, S., Wilikie, B.N. & C.L. Gyles, Resistance of broiler chickens to Escherichia coli respiratory tract infection induced by passively transferred egg-yolk antibodies. Vet. Microbiol. 98: Ketterson, E.D. & V. Nolan, Adaptation, exaptation, and constraint: a hormonal perspective. Am. Nat. 154: Kilpi, M., Hillström, L. & K. Lindström, Egg-size and reproductive success in the Herring Gull Larus argentatus: adaptive or constrained size of the last egg? Ibis 138: Kilner, R., Primary and secondary sexratio manipulation by zebra finches. Anim. Behav. 56: Kitaysky, A.S., Kitaiskaia, E.V., Piatt, J.F. & J.C. Wingfield Benefits and costs of increased levels of corticosterone in seabird chicks. Horm. Behav. 43: Klasing, K.C Nutritional modulation of resistance to infectious diseases. Poultry Sci. 70: Klein, S.L The effects of hormones on sex differences in infection: from genes to behavior. Neurosci. Biobehav. Rev. 24: Komdeur, J., Daan, S., Tinbergen, J. & C. Mateman, Extreme adaptive modification in sex ratio of the Seychelles warbler s eggs. Nature 385: Kowalczyk, K., Daiss, J., Halpern, J. & T.F. Roth, Quantitation of maternalfetal IgG transport in the chicken. Immunology 54:

11 Krijgsveld, K.L., Dijkstra, C. & S. Daan, Energy requirements for growth in relation to sexual size dimorphism in marsh harrier Circus aerogineus nestlings. Physiol. Zool. 71: L Laaksonen, T., Fargallo, J.M., Korpimaki, E., Lyytinen, S., Valkama, J. & V. Poyri, Year- and sex-dependent effects of experimental brood sex ratio manipulations on fledging condition of Eurasian kestrels. J. Anim. Ecol. 73: Lack, D The significance of clutch size. Ibis 89: Legge, S., Heinsohn, R., Double, M.C., Griffiths, R. & A. Cockburn, Complex sex allocation in the laughing kookaburra. Behav. Ecol. 12: Leitner, G., Melamed, D., Drabkin, N. & E.D. Heller, An enzyme-linked immunosorbent assay for detection of antibodies against Escherichia coli: association between hemagglutination test and survival. Avian Dis. 34: Leshchincky, T.V. & K.C. Klasing, Divergence of the inflammatory response in two types of chickens. Develop. Comp. Immunology 25: Leonard, M.L. & P.J. Weatherhead, Dominance and offspring sex ratios in domestic fowl. Anim. Behav. 51: Lessells, C.M. & P.T. Boag, Unrepeatable repeatabilities: a common mistake. Auk 104: Li, Q., Alston-Mills, G.B. & M.A. Ottinger, Avian LHRH-I during embryonic development: Measurement by competitive ELISA with a monoclonal antibody. Gen. Comp. End. 82: Lipar, J.L., Ketterson, E.D. & V. Nolan, Intra-clutch variation in testosterone contents of red-winged blackbirds eggs. Auk 116: Lipar, J.L. & E.D. Ketterson, Maternally derived yolk testosterone enhances the development of the hatching muscle in the red-winged blackbird Aegelius phoeniceus. Proc. Royal Soc. Lond. B 267: Lipar, J.L., Yolk steroids and the development of the hatching muscle in nestling European starlings. J. Avian Biol. 32: Lindström, J Early development and fitness in birds and mammals. TREE 14: Lochmiller, R.L., Vestey, M.R. & J.C. Boren, Relationship between protein nutritional status and immuno-competence in Northern bobwhite chicks. Auk 110: Lochmiller, R.L. & C. Deerenberg, Trade-offs in evolutionary immunology: just what is the cost of immunity? Oikos 88: Love, O.P., Bird, D.M. & L.J. Shutt, Plasma corticosterone in American kestrel siblings: effects of age, hatching order, and hatching asynchrony. Horm. Behav. 43: References 181

12 Loye, J.E. & M. Zuk, Bird-parasite interactions. Oxford University Press, Oxford. M MacRoberts, M.H. & B.R. MacRoberts, The relationship between laying date and incubation period in Herring and lesser black-backed gulls. Ibis 121: Magrath, R.D Hatching asynchrony and reproductive success in the blackbird. Nature 339: Magrath, M.J.L., Brouwer, L. & J. Komdeur, Egg size and laying order in relation to offspring sex in the extreme sexually size dimorphic brown songlark, Cinclorhamphus cruralis. Behav. Ecol. Sociobiol. 54: Marsh, J.A Neuroendocrine-immune interactions in the avian species - a review. Poultry Sci. Rev. 4: Martin II, L.B., Scheuerlein, A. & M. Wikelski, Immune activity elevates energy expenditure of house sparrows: a link between direct and indirect costs. Proc. R. Soc. Lond. B 270: Martinez, J., Potti, J., Davila, J.A. & G. Blanco, Sex, immune response and hatching asynchrony in Mediterranean kestrels (Falco tinnunculus). Abstr. 4 th Eurasian congress on raptors, Seville Martinez, J., Tomas, G., Merino, S., Arriero, E. & J. Moreno, Detection of serum immunoglobulins in wild birds by direct Elisa: a methodological study to validate the technique in different species using antichicken antibodies. Funct. Ecol. 17: Mazuc, J., Bonneaud, C., Chastel, O. & G. Sorci, Social environment affects female and egg testosterone in the house sparrow (Passer domsticus). Ecolgy Letters 6: McGraw K.J. & D.R. Ardia Carotenoids, immunocompetence, and the information content of sexual colors: An experimental test. Am. Nat. 162: Mead, P.S. & M.L. Morton, Hatching asynchrony in the mountain whitecrowned sparrow (Zonotrichia leucophrys oriantha): a selected or incidental trait? Auk 102: Meathrel, C.E Variation in eggs and the period of rapid yolk deposition of the silver gull Larus Novaehollandiae during a protracted laying season. J Zool. 223: Meijer, T., Daan, S. & M. Hall, Family planning in the kestrel (Falco tinnunculus): the proximate control of co-variation of laying date and clutch size. Behaviour 114: Merino, S., Møller, A.P. & F. de Lope, Seasonal changes in cell-mediated immunocompetence and mass gain in nestling barn swallows: a parasitemediated effect. Oikos 90:

13 Metcalfe, N.B. & P. Monaghan, Compensation for a bad start: Grow now pay later? TREE 16: Metcalfe, N.B. & P. Monaghan, Growth versus lifespan: perspectives from evolutionary ecology. Exp. Gerontol. 38: Mock, D.W., Drummond, H. & C.H. Stinson, Avian siblicide. Am. Sci. 78: Monaghan, P. & R. Nager, Why don t lay birds more eggs? TREE 12: Monaghan, P. Nager, R.G. & D.C. Houston, The price of eggs: increased investment in egg production reduces the offspring rearing capacity of parents. Proc. R. Soc. Lond. B 265: Montevecchi, W.A Nest site selection and its survival value among laughing gulls. Behav. Ecol. Sociobiol. 4: Mousseau, T.A. & C.W.Fox, (eds) Maternal effects. New York: Oxford University Press. Müller, W., Dijkstra, C. & T.G.G. Groothuis, Inter-sexual differences in T- cell-mediated immunity of black-headed gull chicks (Larus ridibundus) depend on the hatching order. Behav. Ecol. Sociobiol. 55: Müller W., Eising, C., Dijkstra C. & T.G.G. Groothuis, Sex differences in yolk hormones depend on maternal social status in Leghorn chickens (Gallus gallus domesticus). Proc. R. Soc. Lond. B. 269: Müller, W., Eising, C.M., Dijkstra, C. & T.G.G. Groothuis. Within-clutch patterns of yolk testosterone vary with the onset of incubation in black-headed gulls. Behav. Ecol. in press Müller, W., Groothuis, T.G.G., Dijkstra, C., Siitari, H. & R.V. Alatalo. Maternal antibody transmission and breeding density in the black-headed gull (Larus ridibundus). Funct. Ecol. In press. N Nager, R.G., Monaghan, P., Griffiths, R., Houston, D.C. & R. Dawson, Experimental evidence that off spring sex ratio varies with maternal condition. Proc. Natl. Acad. Sci. USA, 96: Nager, R.G., Monaghan, P., Houston, D.C. & M. Genovart, Parental condition, brood sex ratio and differential young survival: an experimental study in gulls (Larus fuscus). Behav. Ecol. Sociobiol. 48: Nager, R.G., Monaghan, P. & D.C. Houston, 2000b. Within-clutch trade-offs between the number and quality of eggs: experimental manipulations in gulls. Ecology 81: Neuhoff, V., Arold, N., Taube, D & W.Ehrhardt, Improved staining of proteins in polyacrylamide gels including isoelectric focusing gels with clear References 183

14 background at nanogram sensitivity using Coomassie Brilliant Blue G-250 and R-250. Electrophoresis 9: Nielsen, H.C. & J.S. Torday, Sex differences in avian embryo pulmonary surfactant production: evidence for sex chromosome involvement. Endocrinology, 117: Norris, K. & M.R. Evans, Ecological immunology: life history trade-offs and immune defense in birds. Behav. Ecol. 11: O O Connor, R.J Brood reduction in birds: selection for fratricide, infanticide and suicide? Anim. Behav. 26: Ohlsson, T., Smith, H.G., Raberg, L. & D. Hasselquist, Pheasant sexual ornaments reflect nutritional conditions during early growth. Proc. R. Soc. Lond. B 269: Ohlsson, T. & H.G. Smith, Early nutrition causes permanent effects on pheasant morphology. Physiol. Biochem. Zool. 74: Olsen, N.J. & W.J. Kovacs, Gonadal steroids and Immunity. End. Rev. 17: Ots, I. & P. Hõrak, Health impact of blood parasites in breeding great tits. Oecologia 116: Ottinger, M.A. & M.A. Abelnabi, Neuroendocrine system and avian sexual differentiation. Am. Zool. 37: Ottinger, M.A., Pitts, S. & M.A. Abelnabi, Steroid hormones during embryonic development in Japanese Quail: plasma, gonadal, and adrenal levels. Poultry Sci. 80: P Parsons, J., Egg size, laying date and incubation period in the herring gull, Larus argentatus. Ibis 117: 114, Parsons, J., Relationship between egg size and post-hatching chick mortality in the herring gull (Larus argentatus). Nature 228: Parsons, J. 1975a. Asynchronous hatching and chick mortality in the herring gull Larus argentatus. Ibis 117: Parsons, J Factors determining the number and size of eggs laid by the herring gull. Condor 78: Peters, A Testosterone treatment is immunosuppressive in superb fairywrens, yet free-living males with high testosterone are more immunocompetent. Proc. R. Soc. Lond. B 267: Petrie, M., Schwabl, H., Brande-Lavridsen, N. & T. Burke, Sex differences in avian yolk hormone levels. Nature 412: 498. Pike, T.W. & M. Petrie, Potenetial mechanims of avian sex manipulation. Biol. Rev. 78:

15 Pilz, K.M. & H.G. Smith, Egg yolk androgen levels increase with breeding density in the European starling, Sturnus vulgaris. Funct. Ecol. 18: Potti, J. & S. Merino, Parasites and the ontogeny of sexual size dimorphism in a passerine bird. Proc. R. Soc. Lond. B 263: Poxton, I.R Antibodies to lipopolysaccharide. J. Immunol. Meth. 186: Prati, M., Calvo, R. & G. Morreale de Escobar, L-thyroxine and 3,5,3 - triodothyronine concentrations in the chicken egg and in the embryo before and after the onset of thyroid function. Endocrinology 130: R Rasbash, J., Browne, W., Healy, M., Cameron, B. & C. Charlton, Multilevel models project. University of London, London. Reed, W.L. & C.M. Vleck, Functional significance of variation in egg-yolk androgens in the American coot. Oecologia 128: Reid, W.V Constraints on clutch size in the glaucous-winged gull. Stud. Avian Biol. 10: Richner, H The growth dynamics of sexually dimorphic birds and Fisher s sex ratio theory: does sex-specific growth contribute to balanced sex ratios? Funct. Ecol. 5: Richner, H., Oppliger, A. & P. Christe, Effect of an ectoparasite on reproduction in great tits. J. Anim. Ecol. 62: Ricklefs, R.E Comparative analysis of avian embryonic growth. J. Exp. Zool. Suppl. 1: Riedstra, B., Dijkstra, C. & S. Daan,1998. Daily energy expenditure of male and female marsh harrier nestlings. Auk 115: Røskraft, E. & T. Slagsvold, Differential mortality of male and female offspring in experimentally manipulated broods of the rook. J. Anim. Ecol. 54: Ros, A.F.H., Dieleman, S.J. & T.G.G. Groothuis, Social stimuli, testosterone, and aggression in gull chicks: Support for the challenge hypothesis. Horm. Behav. 41: Ros, A.F.H., Groothuis, T.G.G. & V. Apanius, The relation among gonadal steroids, immunocompetence, body mass and behavior in young blackheaded gulls (Larus ridibundus). Am. Nat. 150: Royle, N.J. & K.C. Hamer, Hatching asynchrony and sibling size hierarchies in gulls: effects on parental investment decisions, brood reduction and reproductive success. J. Avian Biol. 29: Royle, N.J., Surai, P.F., McCartney, R.J. & B.K. Speake, Parental investment and egg yolk lipid composition in gulls. Funct. Ecol. 13: Royle, N.J Overproduction in the lesser black-backed gull can marginal chicks overcome the initial handicap of hatching asynchrony? J. Avian Biol. 31: References 185

16 Royle, N.J., Surai, P.F. & I.R. Hartley, Maternal derived androgens and antioxidants in bird eggs: complementary but opposing effects? Behav. Ecol. 12: Rutkovska, J. & M. Cichon, Maternal investment during egg laying and offspring sex: an experimental study of zebra finches. Anim. Behav. 64: S Saino, N., Calza, S. & A.P. Møller, Immunocompetence of nestling barn swallows in relation to brood size and parental effort. J. Anim. Ecol. 66: Saino, N., Martinelli, R. & A.P. Møller, Immunoglobulin plasma concentration in relation to egg laying date and mate ornamentation of female barn swallows (Hirundo rustica). J. Evol. Biol. 14: Saino, N., Romano, M., Ferrari, R.P., Martinelli, R. & A.P. Møller, Maternal antibodies but not carotenoids in barn swallow eggs covary with embryo sex. J. Evol. Biol. 16, Saino, N., Ferrari, R., Romano, M., Martinelli, R. & A.P. Møller, Experimental manipulation of egg carotenoids affects immunity of barn swallow nestlings. Proc. R. Soc. Lond. B 270: Sayag, N., Snapir, N., Robinzon, B., Arnon, E., El Halawani, M.E. & V.E. Grimm, Embryonic sex steroids affect mating behavior and plasma LH in adult chicken. Physiol.& Behav. 45: Sayce, J.R. & G.L. Hunt, Sex ratios of prefledging western gulls. Auk, 104, Schumacher, M., Sulon, J. & J. Balthazart, Changes in serum concentrations of steroids during embryonic and post-hatching development of male and female Japanese quail (Coturnix coturnix japonica). J. Endocrinol.. 118: Schuurs, A.H.W.M. & H.A.M. Verheul, Effects of gender and sex steroids on the immune response. J. Steroid Biochem. 35: Schwabl, H. & E. Kriner, Territorial agression and song of amle robins (Erithacus rubecula) in autumn and spring: Effects of antiandrogen treatment. Horm. Behav. 25: Schwabl, H Yolk is a source of maternal testosterone for developing birds. Proc. Natl. Acad. Sci. USA 90: Schwabl, H. 1996a. Environment modifies the testosterone levels of a female bird and its eggs. J. Exp. Biol. 276: Schwabl, H Maternal testosterone in the egg enhances postnatal growth. Comp. Biochem. Physiol. 114: Schwabl, H., Mock, D.W. & J.A. Gieg, A hormonal mechanism for parental favouritism. Nature 386:

17 Sharp, P.J., Scanes, C.G., Williams, J.B., Harvey, S. & A. Chadwick, Variations in concentrations of prolactin, luteinizing hormone, growth hormone and progesterone in the plasma of broody bantams (Gallus domesticus). J. Endocrinol. 80: Sheldon, B.C. & S. Verhulst, Ecological immunology: costly parasite defenses and trade-offs in evolutionary ecology. TREE 11: Silverin, B Territorial behaviour and hormones of pied flycatchers in optimal and suboptimal habitats. Anim. Behav. 56: Slagsvold, T Hatching asynchrony: interspecific comparisons of altricial birds. Am. Nat. 128: Smith, N.C., Wallach, M., Petracacca, M., Braun, R. & J. Eckert, Maternal transfer of antibodies induced by infection with E. maxima partially protects chickens against challenge with E. tenella. Parasitology 109: Smits, J.E., Bortolotti, G.R. & J.L. Tella, Simplifying the phytohemagglutinin skin-testing technique in studies of avian immunocompetence. Funct. Ecol. 13: Snijders T.A.B. & R.J. Bosker, Multilevel Analysis; An introduction to basic and advanced multilevel modeling. London: SAGE Publications. Sockman, K.W., Schwabl, H. & P.J. Sharp, The role of prolactin in the regulation of clutch size and onset of incubation behavior in the American kestrel. Horm. Behav. 38: Sockman, K.W., Schwabl, H. & P.J. Sharp, Regulation of yolk-androgen concentrations by plasma prolactin in the American kestrel. Horm. Behav. 40: Sockman, K.W. & H. Schwabl, Yolk androgens reduce offspring survival. Proc. R. Soc. Lond. B 267: Soler, J.J., de Neve, L., Perez, T., Soler, M. & G. Sorci, Trade-off between immunocompetence and growth in magpies: An experimental study. Proc. R. Soc. Lond. B 270: Soma, K.K., Sullivan, K. & J.Wingfield, Combined aromatase inhibitor and antiandrogen treatment decreases territorial aggresssion in a wild songbird during nonbreeding season. Gen. Comp. End. 115: Staub, N.L. & M. debeer, The role of androgens in female vertebrates. Gen. Comp. End. 108: Stearns, B.C The evolution of Life Histories. Oxford University Press. Stockland, J.N. & T. Amudsen, Initial size hierarchy in broods of the shag: relative significance of egg size and hatching asynchrony. Auk 107: Stoleson, S.H. & S.R. Beissinger, Hatching asynchrony and the onset of incubation in birds, revisited: When is the critical period? In: Power DM (eds) Current Ornithology 12, pp Plenum, New York. Sturkie, P.D Avian Physiology. Springer, New York. References 187

18 Surai, P.F., Vitamin E in avian reproduction. Poultry Avian Biol. Rev. 10: Surai, P.F., Natural antioxidants in avian nutrition and reproduction. Nottingham University Press. Surai, P.F., Speake, B.K., Noble, R.C. & N.H.C. Sparks, Tissue specific antioxidant profiles and susceptibility to lipid peroxidation of the newly hatched chick. Biol. Trace Elem. Res. 68: Surai, P.F. & B.K. Speake, Distribution of carotenoids from the yolk to the tissues of the chick embryo. J. Nutr. Biochem. 9: T Tanabe, Y., Nakamura, T., Fujoika, K. & O. Doi, Prodcution and secretion of sex steroids by the testes the ovary, and the adrenal glands of embryonic and young chickens (Gallus domesticus). Gen. Comp. End. 39: Tanabe, Y., Takashi, Y. & T. Nakamura, Steroid hormone synthesis and secretion by testes, ovary, and adrenals of embryonic and postembryonic ducks. Gen. Comp. Endocrinol. 49: Tanabe, Y., Saito, N. & T. Nakamura, Ontogenetic steroidgenesis by testes, ovary and adrenals of embryonic and postembryonic chickens (Gallus domesticus). Gen. Comp. Endocrinol. 63: Teather, K.L An experimental study of competition for food between male and female nestlings of the red-winged blackbird. Behav. Ecol. Sociobiol. 31: Tella, J.L., Bortolotti, G.R., Forero, M.G. & R.D. Dawson, Environmental and genetic variation in T-cell-mediated immune response of fledging American kestrels. Oecologia 123: Tella, J.L The evolutionary transition to coloniality promotes higher blood parasitism in birds. J. Evol. Biol. 15: Torres, R. & H. Drummond, Female-biased mortality in daughters of a bird with size dimorphism. J. Anim. Ecol. 66: Torres, R. & H. Drummond, Does large size make daughters of bluefooted booby more expensive than sons? J. Anim. Ecol. 68: Trivers, R.L. & D.E. Willard, 1973 Natural selection of parental ability to vary the sex ratio of offspring. Science 179: Tschirren B., Fitze P.S. & H. Richner, Sexual dimorphism in susceptibility to parasites and cell-mediated immunity in great tit nestlings. J. Anim. Ecol. 72:

19 A Adkins-Regan, E., Ottinger, M.A. & J. Park, Maternal transfer of estradiol to egg yolks alters sexual differentiation of avian offspring. J. Exp. Zool. 271: Allee, W.C., Collias, N.E. & C.Z. Lutherman, Modification of the social order in flocks of hens by the injection of testosterone propionate. Physiol. Zool. 12: Alonso-Alvarez, C. & J.L. Tella, Effects of experimental food restriction and body-mass changes on avian T-cell mediated immune response. Can. J. Zool. 79: Anderson, D.J., Reeve, J., Gomez, J.E.M., Weathers, W.W., Huston, S., Cunningham, H.V. & D.M. Bird, Sexual size dimorphism and food requirements of nestling birds. Can. J. Zool. 71: Anderson, D.J., Budde, C., Apanius, V., Gomez, J.E.M., Bird, D.M. & W.W. Weathers, 1993a. Prey size influences female competitive dominance in nestling American Kestrels Falco sparverius. Ecology 74: Anderson, D.J., Reeve, J. & D.M. Bird, Sexually dimorphic eggs, nestling growth and sibling competition in American kestrels Falco sparverius. Funct. Ecol. 11: Andersson, S., Uller, T., Lõhmus, M., & F. Sundström, Effects of yolk testosterone on growth and immunity in a precocial bird. J. Evol. Biol. 17: Apanius, V., Temple, S.A., & M. Bale, Serum-proteins of wild turkey vultures (Cathartes aura). Comp. Biochem. Physiol. B 76: Apanius, V Ontogeny of immune function. In: Starck, J.M., Ricklefs, R.E. (eds): Avian growth and development evolution within the altricial-precocial. Oxford University Press, Oxford, Chapter 8, pp Apanius, V. & I.C.T. Nisbet, Serum immunoglobulin G levels in very old common terns Sterna hirundo. Exp. Gerontol. 38: Appleby, B.M., Petty, S.J., Blakey, J.K., Rainey, P. & D.W. MacDonald, Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)? Proc. R. Soc. Lond. B 264: Arnold, K.E. & R. Griffiths, Sex-specific hatching order, growth rates and fledging success in jackdaws Corvus monedula. J. Avian Biol. 34: B Badyaev, A.V Growing apart: an ontogenetic perspective on the evolution of sexual size dimorphism. TREE 17: Balthazart, J. & E.A. Adkins-Regan, Sexual differentiation of brain and behaviour in birds. In Hormones, Brain and Behavior vol IV. eds. Pfaff, D., Arnold, A., Etgen, A., Fahrbach, S. and Rabin, R., Elsevier, The Netherlands,

20 V Velando, A Experimental manipulation of maternal effort produces differential effects in sons and daughters: implications for adaptive sex ratios in the blue-footed booby. Behav. Ecol. 13: Verboven, N., Monaghan, P., Evans, D.M., Schwabl, H., Evans, N., Whitelaw, C. & R.G. Nager, Maternal condition, yolk androgens and offspring performance: a supplemental feeding experiment in the lesser black-backed gull (Larus fuscus). Proc. R. Soc. Lond B 270: Viñuela, J Adaptation vs. constraint: intra-clutch egg mass variation in birds. J. Anim. Ecol. 66: von Schantz, T., Bensch, S., Grahn, M., Hasselquist, D. & H. Wittzell, Good genes, oxidative stress and condition-dependent sexual signals. Proc. R. Soc. Lond. B 266: W Walsh, P.S., Metzger, D.A. & R. Higuchi, Chelex-100 as a medium for simple extraction of DNA for PCR-based typing from forensic material. Biotechniques 10: Ward P. & A. Zahavi, The importance of certain assemblages of birds as information-centers for food finding. Ibis 115: Watanabe, S. & F. Nagayama, Studies on the serum IgG level in Japanese quail. Jap. Poult. Sci. 16: Webb, D.R Thermal tolerance of avian embryos: a review. Condor 89: Wittingham, L.A. & H. Schwabl, Maternal testosterone in tree swallow eggs varies with female aggression. Anim. Behav., 63: Williams T.D. (1994) Intraspecific variation in egg size and egg composition in birds: effects on offspring fitness. Biol. Rev. 68: Wilson, J.A. & B. Glick, Ontogeny of mating behaviorin the chicken. Am. J. Physiol. 218: Wingfield, J.C., Hegner, R.E., Dufty, A.M. Jr & G.F. Ball, The challenge hypothesis: theoretical implications for patterns of testosterone secretion, mating systems, and breeding strategies. Am. Nat 136: Woods J.E., Simpson R.M. & P.L. Moore, Plasma testosterone levels in the chick embryo. Gen. Comp. End. 27: Wooler, R.D. & J.C. Coulson Factors affecting the age of first breeding on the kittiwake Rissa tridactyla. Ibis 119: References 189

21 190