YU Lei，RAO Kai-qing，DONG Chuan-hao，XU Ya-ou. Programming Effects of Glucocorticoids in Eggs on the Growth of Offspring[J]. ACTA VETERINARIA ET ZOOTECHNICA SINICA, 2015, 46(2): 186-191.
［1］SCHWABL H.Yolk is a source of maternal testosterone for developing birds［J］. Proc Natl Acad Sci U S A，1993，90(24)：11446-11450.
［2］HAYWARD L S，SATTERLEE D G，WINGFIELD J C.Japanese quail selected for high plasma corticosterone response deposit high levels of corticosterone in their eggs［J］.Physiol Biochem Zool，2005，78(6)：1026-1031.
［3］LOVE O P，WILLIAMS T D.The adaptive value of stress-induced phenotypes：effects of maternally derived corticosterone on sex-biased investment，cost of reproduction，and maternal fitness［J］.Am Nat，2008，172(4)：E135-E149.
［4］RUBOLINI D，ROMANO M，BONCORAGLIO G，et al.Effects of elevated egg corticosterone levels on behavior，growth，and immunity of yellow-legged gull (Larus michahellis) chicks［J］.Horm Behav，2005，47(5)：592-605.
［5］RETTENBACHER S，MÖSTL E，HACKL R，et al.Corticosterone in chicken eggs［J］.Ann N Y Acad Sci，2005，1046(1)：193-203.
［6］RETTENBACHER S，MÖSTL E，GROOTHUIS T G G.Gestagens and glucocorticoids in chicken eggs［J］.Gen Comp Endocrinol，2009，164(2)：125-129.
［7］NAVARA K J，PINSON S E.Yolk and albumen corticosterone concentrations in eggs laid by white versus brown caged laying hens［J］. Poult Sci，2010，89(7)：1509-1513.
［8］BOWDEN R M，EWERT M A，LIPAR J L，et al.Concentrations of steroid hormones in layers and biopsies of chelonian egg yolks［J］.Gen Comp Endocrinol，2001，121(1)：95-103.
［9］ROYO F，MAYO S，CARLSSON H E，et al.Egg corticosterone：A noninvasive measure of stress in egg-laying birds［J］. J Avian Med Surg，2008，22(4)：310-314.
［10］JOHNSON A L.Avian physiology［M］.4th ed .New York：Springer-Verlag，1986：403-431.
［11］DOWNING J A，BRYDEN W L.Determination of corticosterone concentrations in egg albumen：a non-invasive indicator of stress in laying hens［J］. Physiol Behav，2008，95(3)：381-387.
［12］TANABE Y，SAITO N，NAKAMURA T.Ontogenetic steroidogenesis by testes，ovary，and adrenals of embryonic and postembryonic chickens(Gallus domesticus)［J］.Gen Comp Endocrinol，1986，63(3)：456-463.
［13］CARSIA R V，MORIN M E，WEBER H.Ontogenic corticosteroidogenesis of the domestic fowl：response of isolated adrenocortical cells［J］.Proc Soc Exp Biol Med，1987，184(4)：436-445.
［14］MARIE C.Ontogenesis of the adrenal glucocorticoids and of the target function of the enzymatic tyrosine transaminase activity in the chick embryo［J］.J Endocrinol，1981，90(2)：193-200.
［15］OKULIAROVÁ M，ŠÁRNIKOVÁ B，RETTENBA-CHERÁ S，et al.Yolk testosterone and corticosterone in hierarchical follicles and laid eggs of Japanese quail exposed to long-term restraint stress［J］.Gen Comp Endocrinol，2010，165(1)：91-96.
［16］AHMED A A，MA W，GUO F，et al.Differences in egg deposition of corticosterone and embryonic expression of corticosterone metabolic enzymes between slow and fast growing broiler chickens［J］.Comp Biochem Physiol A Mol Integr Physiol，2013，164(1)：200-206.
［17］饶开晴.Leptin 对鸡胚卵黄膜和尿囊膜基因表达及血管形成的影响［D］.南京：南京农业大学，2009.
RAO K Q.Effect of leptin on yolk sac and chorioallantoic membrane gene expression and angiogenesis in the chicken embryo［D］.Nanjing：Nanjing Agricultural University，2009.(in Chinese)
［18］AHMED A A，MA W，NI Y，et al.Corticosterone in ovo modifies aggressive behaviors and reproductive performances through alterations of the hypothalamic-pituitary-gonadal axis in the chicken［J］.Anim Reprod Sci，2014，146(3)：193-201.
［19］KIRSCHBAUM C，WÜST S，FAIG H G，et al.Heritability of cortisol responses to human corticotropin-releasing hormone，ergometry，and psychological stress in humans［J］.J Clin Endocrinol Metab，1992，75(6)：1526-1530.
［20］DÉSAUTÉS C，BIDANEL J P，MORMÈDE P.Genetic study of behavioral and pituitary-adrenocortical reactivity in response to an environmental challenge in pigs［J］.Physiol Behav，1997，62(2)：337-345.
［21］SARRIEAU A，MORMEDE P.Hypothalamic-pituitary-adrenal axis activity in the inbred Brown Norway and Fischer 344 rat strains［J］.Life Sci，1998，62(16)：1417-1425.
［22］李留安，吴展望，赵茹茜.母体妊娠期应激对后代出生前后下丘脑-垂体-肾上腺轴的影响及其机制［J］.动物营养学报，2010，22(5)：1138-1141.
LI L A，WU Z W，ZHAO R Q.Effects of maternal stress during pregnancy on offspring before and after birth：mechanism of action from a Hypothalamus-pituitary-adrenocortical axis［J］.Chinese Journal of Animal Nutrition，2010，22(5)：1138-1141.(in Chinese)
［23］HAYWARD L S，WINGFIELD J C.Maternal corticosterone is transferred to avian yolk and may alter offspring growth and adult phenotype［J］.Gen Comp Endocrinol，2004，135(3)：365-371.
［24］ZIMMER C，BOOGERT N J，SPENCER K A.Developmental programming：Cumulative effects of increased pre-hatching corticosterone levels and post-hatching unpredictable food availability on physiology and behaviour in adulthood［J］.Horm Behav，2013，64(3)：494-500.
［25］MARTINS T L F，ROBERTS M L，GIBLIN I，et al.Speed of exploration and risk-taking behavior are linked to corticosterone titres in zebra finches［J］.Horm Behav，2007，52(4)：445-453.
［26］STÖWE M，ROSIVALL B，DRENT P J，et al.Selection for fast and slow exploration affects baseline and stress-induced corticosterone excretion in Great tit nestlings，Parus major［J］.Horm Behav，2010，58(5)：864-871.
［27］CATALANI A，CASOLINI P，SCACCIANOCE S，et al.Maternal corticosterone during lactation permanently affects brain corticosteroid receptors，stress response and behaviour in rat progeny［J］.Neuroscience，2000，100(2)：319-325.
［28］NYIRENDA M J，LINDSAY R S，KENYON C J，et al.Glucocorticoid exposure in late gestation permanently programs rat hepatic phosphoenolpyruvate carboxy kinase and glucocorticoid receptor expression and causes glucose intolerance in adult offspring［J］.J Clin Invest，1998，101(10)：2174.
［29］REYNOLDS L P，BOROWICZ P P，CATON J S，et al.Developmental programming：the concept，large animal models，and the key role of uteroplacental vascular development［J］.J Anim Sci，2010，88(13)：E61-E72.
［30］BALE T L.Neuroendocrine and immune influences on the CNS：It’s a matter of sex［J］.Neuron，2009，64(1)：13-16.
［31］KALLIECHARAN R，HALL B K.A developmental study of the levels of progesterone，corticosterone，cortisol，and cortisone circulating in plasma of chick embryos［J］.Gen Comp Endocrinol，1974，24(4)：364-372.
［32］高静.糖皮质激素诱导肉鸡代谢程序化效应的研究［D］.泰安：山东农业大学，2008.
GAO J.Metabolic programming induced by glucocorticoids in broiler chickens［D］.Taian：Shandong Agricultural University，2008.(in Chinese)
［33］ERIKSEN M S，HAUG A，TORJESEN P A，et al.Prenatal exposure to corticosterone impairs embryonic development and increases fluctuating asymmetry in chickens(Gallus gallus domesticus)［J］.Br Poult Sci，2003，44(5)：690-697.
［34］PIKE T W，PETRIE M.Experimental evidence that corticosterone affects offspring sex ratios in quail［J］. Proc Biol Sci，2006，273(1590)：1093-1098.
［35］HEIBLUM R，ARNON E，CHAZAN G，et al.Glucocorticoid administration during incubation:embryo mortality and posthatch growth in chickens［J］. Poult Sci，2001，80(9)：1357-1363.
［36］高静，宋志刚，焦洪超，等.不同胚龄皮质酮处理对鸡胚胎生长发育的影响［J］.畜牧兽医学报，2008，39(9)：1224-1229.
GAO J，SONG Z G，JIAO H C.Effect of corticosterone treatment at different hatching time on embryo development of broiler chickens［J］.Acta Veterinaria et Zootechnica Sinica，2008，39(9)：1224-1229.(in Chinese)
［37］HEIJMANS B T，TOBI E W，STEIN A D，et al.Persistent epigenetic differences associated with prenatal exposure to famine in humans［J］.Proc Natl Acad Sci U S A，2008，105(44)：17046-17049.
［38］OBERLANDER T F，WEINBERG J，PAPSDORF M，et al.Prenatal exposure to maternal depression，neonatal methylation of human glucocorticoid receptor gene (NR3C1) and infant cortisol stress responses［J］.Epigenetics，2008，3(2)：97-106.
［39］MCGOWAN P O，SASAKI A，D′ALESSIO A C，et al.Epigenetic regulation of the glucocorticoid receptor in human brain associates with childhood abuse［J］.Nat Neurosci，2009，12(3)：342-348.
［40］WEAVER I C，CERVONI N，CHAMPAGNE F A，et al.Epigenetic programming by maternal behavior［J］.Nat Neurosci，2004，7(8)：847-854.
［41］MURGATROYD C，PATCHEV A V，WU Y，et al.Dynamic DNA methylation programs persistent adverse effects of early-life stress［J］.Nat Neurosci，2009，12(12)：1559-1566.
［42］WANG S，NI Y，GUO F，et al.Differential expression of hypothalamic fear and stress-related genes in broiler chickens showing short or long tonic immobility［J］.Domest Anim Endocrinol，2014，47：65-72.
