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T�I	kTT�V�V�,�������C������fM>�VC`:n<vn�V,�s�O��s�V�s��VX	1=N@\T�B��D�1=1=1=/k/kjQ�1=1=1=vn�I�I�I�I���������������������������������������������������������������������s1=1=1=1=1=1=1=1=1=\ |:	Prenatal corticosterone exposure alters glucocorticoid metabolic enzyme gene mRNA associated with increased aggressive behaviors and tonic immobility in chicken
Abdelkareem A. Ahmeda,b,c,d,*, Mohammed Elmujtba Adam Essad, Adriano Mollicad,e, Azzurra Stefanuccid,e, Gokhan Zengind,f, Hussein Ahmedg, Ayman Sati Sati Mohamedd
a Department of Physiology and Biochemistry, Faculty of Veterinary Science, University of Nyala, Nyala, Sudan
  b Institute of Molecular Biology, University of Nyala, Nyala, Sudan 
c Biomedical Research Institute, Darfur University College, Nyala, Sudan
d Department of Clinical Medicine, Medical and cancer Research Institute, Nyala, Sudan
e Department of Pharmacy, University �G. d�Annunzio� of Chieti-Pescara, 66100, Chieti, Italy
fDepartment of Biology, Science Faculty, Selcuk University, Konya, Turkey
g College of Veterinary and Animal Science, the Islamia University, Bahawalpur, Pakistan
*Corresponding author: 
Dr. Abdelkareem Abdallah Ahmed 
Institute of Molecular Biology,University of Nyala, Nyala, Sudan
Email:� HYPERLINK "mailto:kareemo151@gmail.com" \t "_blank" kareemo151@gmail.com,  HYPERLINK "mailto:kareemo151@vet.nyalau.edu.sd" \t "_blank" kareemo151@vet.nyalau.edu.sd, P.O Box: 155 Nyala, Sudan
Tel: +249909998350, +249923433164
Fax: 00249711833123 

Abstract: Exposure to excess glucocorticoids (GCs) during embryonic development influences offspring physiology and behaviors and induces change in hypothalamic-pituitary-adrenal (HPA) axis genes expression and serotonergic system in mammals. Whether prenatal corticosterone (CORT) exposure induces similar effects in avian species remains unclear. In the present study, we injected low (0.2 �g) and high (1 �g) doses of CORT in ovo before incubation and detected changes in aggressive behavior, tonic immobility (TI), HPA axis and 5-hydroxytryptamine (serotonin) (5-HT) system gene expression on posthatch chickens of different ages. High dose of CORT significantly (P<0.05) suppressed growth rate, increased the frequency of aggressive behaviors, which was associated with elevated plasma CORT concentration. Likewise, in ovo injection of CORT significantly (P<0.05) increased tonic immobility (TI) duration both in chickens from low and high doses of CORT treatments compared to control. In addition, administration of CORT significantly (P<0.05) up-regulated mRNA expression of 11�-hydroxysteroid dehydrogenase type 1 (11�-HSD1) whereas it down-regulated 11�-hydroxysteroid dehydrogenase type 2 (11�-HSD2) and mineralocorticoid receptor (MR) mRNA expression in the hypothalamus. No significant differences were seen in glucocorticoid receptor (GR) and 20-hydroxysteroid dehydrogenase (20-HSD) mRNA levels upon CORT treatment. Moreover, CORT exposure significantly (P<0.05) increased hypothalamic 5-hydroxytryptamine (serotonin) receptor 1A (5-HTR1A) mRNA expression, but not 5-HT receptor 1B (5-HTR1B). In ovo administration of CORT may programs the aggressive behaviors in the chicken through alterations of HPA axis and 5-HT system. 
Keywords: behavior, chicken, corticosterone, glucocorticoid metabolic enzymes, hypothalamus 
Introduction
    The phenotype of an individual is not only programmed by genetic factor, however, its also via an ecological factors that play a fundamental role in determining offspring phenotype  ADDIN EN.CITE <EndNote><Cite><Author>Nijhout</Author><Year>2003</Year><RecNum>50529</RecNum><DisplayText>[1]</DisplayText><record><rec-number>50529</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">50529</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Nijhout, H. F.</author></authors></contributors><auth-address>Department of Biology, Duke University, Durham, NC 27708, USA. hfn@duke.edu</auth-address><titles><title>Development and evolution of adaptive polyphenisms</title><secondary-title>Evol Dev</secondary-title></titles><periodical><full-title>Evol Dev</full-title></periodical><pages>9-18</pages><volume>5</volume><number>1</number><edition>2002/12/21</edition><keywords><keyword>*Adaptation, Physiological</keyword><keyword>Animals</keyword><keyword>Beetles</keyword><keyword>*Biological Evolution</keyword><keyword>Ecology</keyword><keyword>Environment</keyword><keyword>Female</keyword><keyword>*Gene Expression Regulation, Developmental</keyword><keyword>Genotype</keyword><keyword>Male</keyword><keyword>Metamorphosis, Biological</keyword><keyword>Phenotype</keyword><keyword>Seasons</keyword></keywords><dates><year>2003</year><pub-dates><date>Jan-Feb</date></pub-dates></dates><isbn>1520-541X (Print)&#xD;1520-541X (Linking)</isbn><accession-num>12492404</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12492404</url></related-urls></urls><electronic-resource-num>ede03003 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_1" \o "Nijhout, 2003 #50529" 1], physiology  ADDIN EN.CITE <EndNote><Cite><Author>Brouwer</Author><Year>2019</Year><RecNum>67216</RecNum><DisplayText>[2]</DisplayText><record><rec-number>67216</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67216</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Brouwer, Lyanne</author><author>Griffith, Simon C.</author></authors></contributors><titles><title>Extra-pair paternity in birds</title><secondary-title>Molecular Ecology</secondary-title></titles><periodical><full-title>Mol Ecol</full-title><abbr-1>Molecular ecology</abbr-1></periodical><pages>4864-4882</pages><volume>28</volume><number>22</number><dates><year>2019</year></dates><isbn>0962-1083</isbn><urls><related-urls><url>https://onlinelibrary.wiley.com/doi/abs/10.1111/mec.15259</url></related-urls></urls><electronic-resource-num>10.1111/mec.15259</electronic-resource-num></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_2" \o "Brouwer, 2019 #67216" 2], and behavior  ADDIN EN.CITE <EndNote><Cite><Author>Kerhoas</Author><Year>2014</Year><RecNum>67215</RecNum><DisplayText>[3]</DisplayText><record><rec-number>67215</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67215</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kerhoas, Daphne</author><author>Perwitasari-Farajallah, Dyah</author><author>Agil, Muhammad</author><author>Widdig, Anja</author><author>Engelhardt, Antje</author></authors></contributors><titles><title>Social and ecological factors influencing offspring survival in wild macaques</title><secondary-title>Behavioral ecology : official journal of the International Society for Behavioral Ecology</secondary-title><alt-title>Behav Ecol</alt-title></titles><periodical><full-title>Behavioral ecology : official journal of the International Society for Behavioral Ecology</full-title><abbr-1>Behav Ecol</abbr-1></periodical><alt-periodical><full-title>Behavioral ecology : official journal of the International Society for Behavioral Ecology</full-title><abbr-1>Behav Ecol</abbr-1></alt-periodical><pages>1164-1172</pages><volume>25</volume><number>5</number><edition>06/17</edition><keywords><keyword>Macaca nigra</keyword><keyword>between-group encounters</keyword><keyword>female reproductive success</keyword><keyword>offspring loss</keyword><keyword>proportional hazards model</keyword><keyword>socioecology.</keyword></keywords><dates><year>2014</year></dates><publisher>Oxford University Press</publisher><isbn>1045-2249&#xD;1465-7279</isbn><accession-num>25214754</accession-num><urls><related-urls><url>https://pubmed.ncbi.nlm.nih.gov/25214754</url><url>https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4160111/</url></related-urls></urls><electronic-resource-num>10.1093/beheco/aru099</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_3" \o "Kerhoas, 2014 #67215" 3].  In birds, maternal influences have aroused much interest after the discovery that bird's eggs contain a variety of maternal derived steroid hormones  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_4" \o "Groothuis, 2005 #2309" 4,  HYPERLINK \l "_ENREF_5" \o "Schwabl, 1993 #1764" 5]. Corticosterone (CORT), the main plasma glucocorticoid (GC) in avian species has been confirmed to be transmitted into chickens egg  ADDIN EN.CITE <EndNote><Cite><Author>Rettenbacher</Author><Year>2009</Year><RecNum>1250</RecNum><DisplayText>[6]</DisplayText><record><rec-number>1250</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">1250</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rettenbacher, S.</author><author>Mostl, E.</author><author>Groothuis, T. G.</author></authors></contributors><auth-address>Department of Biomedical Sciences, Biochemistry, University of Veterinary Medicine Vienna, Veterinarplatz 1, 1210 Vienna, Austria. sophie.rettenbacher@vu-wien.ac.at</auth-address><titles><title>Gestagens and glucocorticoids in chicken eggs</title><secondary-title>Gen Comp Endocrinol</secondary-title></titles><periodical><full-title>Gen Comp Endocrinol</full-title></periodical><pages>125-9</pages><volume>164</volume><number>2-3</number><edition>2009/06/09</edition><keywords><keyword>Animals</keyword><keyword>*Chickens</keyword><keyword>Chromatography, High Pressure Liquid</keyword><keyword>Corticosterone/analysis</keyword><keyword>Egg Yolk/chemistry</keyword><keyword>Eggs/*analysis</keyword><keyword>Female</keyword><keyword>Glucocorticoids/*analysis/immunology</keyword><keyword>Pregnanes/analysis</keyword><keyword>Pregnenolone/analysis</keyword><keyword>Progesterone/analysis</keyword><keyword>Progestins/*analysis/immunology</keyword></keywords><dates><year>2009</year><pub-dates><date>Nov-Dec</date></pub-dates></dates><isbn>1095-6840 (Electronic)&#xD;0016-6480 (Linking)</isbn><accession-num>19501091</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=19501091</url></related-urls></urls><electronic-resource-num>S0016-6480(09)00227-5 [pii]&#xD;10.1016/j.ygcen.2009.05.019</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_6" \o "Rettenbacher, 2009 #1250" 6]. The CORT concentration in eggs has reported to be modified via a variety of factors including physiological status of the bird  ADDIN EN.CITE <EndNote><Cite><Author>Saino</Author><Year>2005</Year><RecNum>19894</RecNum><DisplayText>[7]</DisplayText><record><rec-number>19894</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">19894</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Saino, N.</author><author>Romano, M.</author><author>Ferrari, R. P.</author><author>Martinelli, R.</author><author>Moller, A. P.</author></authors></contributors><auth-address>Dipartimento di Biologia, Universita degli Studi di Milano, via Celoria 26, I-20133 Milano, Italy. nicola.saino@unimi.it</auth-address><titles><title>Stressed mothers lay eggs with high corticosterone levels which produce low-quality offspring</title><secondary-title>J Exp Zool A Comp Exp Biol</secondary-title></titles><periodical><full-title>J Exp Zool A Comp Exp Biol</full-title></periodical><pages>998-1006</pages><volume>303</volume><number>11</number><edition>2005/10/12</edition><keywords><keyword>Analysis of Variance</keyword><keyword>Animals</keyword><keyword>Animals, Newborn/*growth &amp; development</keyword><keyword>Bird Diseases/*metabolism/*physiopathology</keyword><keyword>Corticosterone/*metabolism/toxicity</keyword><keyword>Female</keyword><keyword>Italy</keyword><keyword>Ovum/drug effects/*metabolism</keyword><keyword>Stress, Physiological/metabolism/physiopathology/*veterinary</keyword><keyword>*Swallows</keyword></keywords><dates><year>2005</year><pub-dates><date>Nov 1</date></pub-dates></dates><isbn>1548-8969 (Print)&#xD;1548-8969 (Linking)</isbn><accession-num>16217808</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=16217808</url></related-urls></urls><electronic-resource-num>10.1002/jez.a.224</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_7" \o "Saino, 2005 #19894" 7], stressful surroundings  environment  ADDIN EN.CITE <EndNote><Cite><Author>Hayward</Author><Year>2004</Year><RecNum>1602</RecNum><DisplayText>[8]</DisplayText><record><rec-number>1602</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">1602</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hayward, L. S.</author><author>Wingfield, J. C.</author></authors></contributors><auth-address>Department of Zoology, University of Washington, Seattle, WA 98195, USA. lhayward@u.washington.edu</auth-address><titles><title>Maternal corticosterone is transferred to avian yolk and may alter offspring growth and adult phenotype</title><secondary-title>Gen Comp Endocrinol</secondary-title></titles><periodical><full-title>Gen Comp Endocrinol</full-title></periodical><pages>365-71</pages><volume>135</volume><number>3</number><edition>2004/01/16</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn</keyword><keyword>Birth Weight/drug effects</keyword><keyword>Corticosterone/blood/metabolism/*pharmacology</keyword><keyword>Coturnix/growth &amp; development/*metabolism</keyword><keyword>Egg Yolk/chemistry/*metabolism</keyword><keyword>Female</keyword><keyword>Male</keyword><keyword>Reproduction/drug effects</keyword><keyword>Stress, Physiological/metabolism</keyword><keyword>Zygote/chemistry/metabolism</keyword></keywords><dates><year>2004</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0016-6480 (Print)&#xD;0016-6480 (Linking)</isbn><accession-num>14723888</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=14723888</url></related-urls></urls><electronic-resource-num>S0016648003004039 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_8" \o "Hayward, 2004 #1602" 8], and housing system  ADDIN EN.CITE <EndNote><Cite><Author>Lay</Author><Year>2011</Year><RecNum>32149</RecNum><DisplayText>[9]</DisplayText><record><rec-number>32149</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">32149</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Lay, D. C., Jr.</author><author>Fulton, R. M.</author><author>Hester, P. Y.</author><author>Karcher, D. M.</author><author>Kjaer, J. B.</author><author>Mench, J. A.</author><author>Mullens, B. A.</author><author>Newberry, R. C.</author><author>Nicol, C. J.</author><author>O&apos;Sullivan, N. P.</author><author>Porter, R. E.</author></authors></contributors><auth-address>Livestock Behavior Research Unit, Agricultural Research Service-USDA, West Lafayette, IN 47907, USA. Don.Lay@ars.usda.gov</auth-address><titles><title>Hen welfare in different housing systems</title><secondary-title>Poult Sci</secondary-title></titles><periodical><full-title>Poult Sci</full-title></periodical><pages>278-94</pages><volume>90</volume><number>1</number><edition>2010/12/24</edition><keywords><keyword>Animal Welfare/*standards</keyword><keyword>Animals</keyword><keyword>Chickens/*physiology</keyword><keyword>Eggs/microbiology</keyword><keyword>Female</keyword><keyword>Housing, Animal/*standards</keyword><keyword>Poultry Diseases/prevention &amp; control</keyword></keywords><dates><year>2011</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0032-5791 (Print)&#xD;0032-5791 (Linking)</isbn><accession-num>21177469</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=21177469</url></related-urls></urls><electronic-resource-num>90/1/278 [pii]&#xD;10.3382/ps.2010-00962</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_9" \o "Lay, 2011 #32149" 9].
    Fetuses or embryos of mammals and avian species are exposed to a substantial amount of maternal GCs either through the placenta in mammals  ADDIN EN.CITE <EndNote><Cite><Author>Seckl</Author><Year>2004</Year><RecNum>17842</RecNum><DisplayText>[10]</DisplayText><record><rec-number>17842</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">17842</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Seckl, J. R.</author></authors></contributors><auth-address>Endocrinology Unit, University of Edinburgh, Western General Hospital, Edinburgh EH4 2XU, UK. J.Seckl@ed.ac.uk</auth-address><titles><title>Prenatal glucocorticoids and long-term programming</title><secondary-title>Eur J Endocrinol</secondary-title></titles><periodical><full-title>Eur J Endocrinol</full-title></periodical><pages>U49-62</pages><volume>151 Suppl 3</volume><edition>2004/11/24</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn/embryology</keyword><keyword>Behavior</keyword><keyword>Diabetes Mellitus, Type 1/etiology</keyword><keyword>Female</keyword><keyword>Glucocorticoids/*adverse effects/physiology</keyword><keyword>Humans</keyword><keyword>Hydrocortisone/blood</keyword><keyword>Hypertension/etiology</keyword><keyword>Infant, Low Birth Weight/blood/growth &amp; development</keyword><keyword>Infant, Newborn</keyword><keyword>Lung/embryology</keyword><keyword>Pregnancy</keyword><keyword>*Prenatal Exposure Delayed Effects</keyword><keyword>Rats</keyword><keyword>Stress, Physiological/complications</keyword></keywords><dates><year>2004</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>0804-4643 (Print)&#xD;0804-4643 (Linking)</isbn><accession-num>15554887</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15554887</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_10" \o "Seckl, 2004 #17842" 10] or by yolk deposition in birds  ADDIN EN.CITE <EndNote><Cite><Author>Saino</Author><Year>2005</Year><RecNum>2476</RecNum><DisplayText>[7]</DisplayText><record><rec-number>2476</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">2476</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Saino, N.</author><author>Romano, M.</author><author>Ferrari, R. P.</author><author>Martinelli, R.</author><author>Moller, A. P.</author></authors></contributors><auth-address>Dipartimento di Biologia, Universita degli Studi di Milano, via Celoria 26, I-20133 Milano, Italy. nicola.saino@unimi.it</auth-address><titles><title>Stressed mothers lay eggs with high corticosterone levels which produce low-quality offspring</title><secondary-title>J Exp Zool A Comp Exp Biol</secondary-title></titles><periodical><full-title>J Exp Zool A Comp Exp Biol</full-title></periodical><pages>998-1006</pages><volume>303</volume><number>11</number><edition>2005/10/12</edition><keywords><keyword>Analysis of Variance</keyword><keyword>Animals</keyword><keyword>Animals, Newborn/*growth &amp; development</keyword><keyword>Bird Diseases/*metabolism/*physiopathology</keyword><keyword>Corticosterone/*metabolism/toxicity</keyword><keyword>Female</keyword><keyword>Italy</keyword><keyword>Ovum/drug effects/*metabolism</keyword><keyword>Stress, Physiological/metabolism/physiopathology/*veterinary</keyword><keyword>*Swallows</keyword></keywords><dates><year>2005</year><pub-dates><date>Nov 1</date></pub-dates></dates><isbn>1548-8969 (Print)&#xD;1548-8969 (Linking)</isbn><accession-num>16217808</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=16217808</url></related-urls></urls><electronic-resource-num>10.1002/jez.a.224</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_7" \o "Saino, 2005 #19894" 7]. Chronic stress modulates the hypothalamic-pituitary-adrenal (HPA) axis function results in increased exposure to GCs via elevation in baseline of GCs levels and thus causes a decrease in offspring body weight gain and growth  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_11" \o "Eriksen, 2003 #7189" 11,  HYPERLINK \l "_ENREF_12" \o "Rubolini, 2005 #1559" 12]. Exposure of GCs during the development of embryos is confirmed to have both short- and long-term outcomes  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_10" \o "Seckl, 2004 #17842" 10,  HYPERLINK \l "_ENREF_13" \o "Love, 2008 #2377" 13], for example it reduced offspring weight  ADDIN EN.CITE <EndNote><Cite><Author>Janczak</Author><Year>2006</Year><RecNum>37868</RecNum><DisplayText>[14]</DisplayText><record><rec-number>37868</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37868</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Janczak, A. M.</author><author>Braastad, B. O.</author><author>Bakken, M.</author></authors></contributors><auth-address>Janczak, AM&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, N-1432 As, Norway</auth-address><titles><title>Behavioural effects of embryonic exposure to corticosterone in chickens</title><secondary-title>Applied Animal Behaviour Science</secondary-title><alt-title>Appl Anim Behav Sci</alt-title></titles><periodical><full-title>Applied Animal Behaviour Science</full-title></periodical><alt-periodical><full-title>Appl Anim Behav Sci</full-title></alt-periodical><pages>69-82</pages><volume>96</volume><number>1-2</number><keywords><keyword>prenatal stress</keyword><keyword>chicken</keyword><keyword>fear</keyword><keyword>cognition</keyword><keyword>competition</keyword><keyword>prenatal stress</keyword><keyword>feather pecking</keyword><keyword>plasma-corticosterone</keyword><keyword>pregnancy</keyword><keyword>animals</keyword><keyword>anxiety</keyword><keyword>alters</keyword><keyword>yolk</keyword><keyword>rats</keyword><keyword>hen</keyword></keywords><dates><year>2006</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0168-1591</isbn><accession-num>ISI:000235262600006</accession-num><urls><related-urls><url>&lt;Go to ISI&gt;://000235262600006</url></related-urls></urls><electronic-resource-num>DOI 10.1016/j.applanim.2005.04.020</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_14" \o "Janczak, 2006 #37863" 14] and compromised immune system function  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_12" \o "Rubolini, 2005 #1559" 12]. Prenatal exposure of GC modify the HPA axis activity  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_15" \o "Moisiadis, 2017 #67217" 15,  HYPERLINK \l "_ENREF_16" \o "Maniam, 2014 #67239" 16] and behavior  ADDIN EN.CITE <EndNote><Cite><Author>Seckl</Author><Year>2004</Year><RecNum>8103</RecNum><DisplayText>[17]</DisplayText><record><rec-number>8103</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">8103</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Seckl, J. R.</author><author>Meaney, M. J.</author></authors></contributors><auth-address>Endocrinology Unit, University of Edinburgh, Molecular Medicine Centre, Western General Hospital, Edinburgh, EH4 2XU, UK. J.Seckl@ed.ac.uk</auth-address><titles><title>Glucocorticoid programming</title><secondary-title>Ann N Y Acad Sci</secondary-title></titles><periodical><full-title>Ann N Y Acad Sci</full-title></periodical><pages>63-84</pages><volume>1032</volume><edition>2005/01/29</edition><keywords><keyword>11-beta-Hydroxysteroid Dehydrogenase Type 2/metabolism</keyword><keyword>Behavior/physiology</keyword><keyword>Birth Weight/physiology</keyword><keyword>Cardiovascular Physiological Phenomena</keyword><keyword>Central Nervous System/embryology/physiology</keyword><keyword>Female</keyword><keyword>Fetal Development/physiology</keyword><keyword>Glucocorticoids/*physiology</keyword><keyword>Humans</keyword><keyword>Infant, Newborn</keyword><keyword>Infant, Premature</keyword><keyword>Metabolism/physiology</keyword><keyword>Pregnancy</keyword><keyword>Prenatal Exposure Delayed Effects</keyword><keyword>Stress, Physiological/physiopathology</keyword></keywords><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>0077-8923 (Print)&#xD;0077-8923 (Linking)</isbn><accession-num>15677396</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15677396</url></related-urls></urls><electronic-resource-num>1032/1/63 [pii]&#xD;10.1196/annals.1314.006</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_17" \o "Seckl, 2004 #8103" 17]. In addition, In ovo injection of CORT prior to incubation has found to increase flight performance behavior  ADDIN EN.CITE <EndNote><Cite><Author>Chin</Author><Year>2009</Year><RecNum>37318</RecNum><DisplayText>[18]</DisplayText><record><rec-number>37318</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37318</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Chin, E. H.</author><author>Love, O. P.</author><author>Verspoor, J. J.</author><author>Williams, T. D.</author><author>Rowley, K.</author><author>Burness, G.</author></authors></contributors><auth-address>Environmental and Life Sciences Graduate Program, Trent University, Peterborough, Ontario, Canada K9J 7B8.</auth-address><titles><title>Juveniles exposed to embryonic corticosterone have enhanced flight performance</title><secondary-title>Proc Biol Sci</secondary-title></titles><periodical><full-title>Proc Biol Sci</full-title></periodical><pages>499-505</pages><volume>276</volume><number>1656</number><edition>2008/10/10</edition><keywords><keyword>Animals</keyword><keyword>*Corticosterone</keyword><keyword>Embryo, Nonmammalian</keyword><keyword>*Flight, Animal</keyword><keyword>Starlings/*physiology</keyword><keyword>Stress, Physiological</keyword></keywords><dates><year>2009</year><pub-dates><date>Feb 7</date></pub-dates></dates><isbn>0962-8452 (Print)&#xD;0962-8452 (Linking)</isbn><accession-num>18842541</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=18842541</url></related-urls></urls><custom2>2664354</custom2><electronic-resource-num>F6H80G1662207650 [pii]&#xD;10.1098/rspb.2008.1294</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_18" \o "Chin, 2009 #37318" 18], fearfulness behavior  ADDIN EN.CITE <EndNote><Cite><Author>Janczak</Author><Year>2006</Year><RecNum>37863</RecNum><DisplayText>[14]</DisplayText><record><rec-number>37863</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37863</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Janczak, A. M.</author><author>Braastad, B. O.</author><author>Bakken, M.</author></authors></contributors><auth-address>Janczak, AM&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, N-1432 As, Norway</auth-address><titles><title>Behavioural effects of embryonic exposure to corticosterone in chickens</title><secondary-title>Applied Animal Behaviour Science</secondary-title><alt-title>Appl Anim Behav Sci</alt-title></titles><periodical><full-title>Applied Animal Behaviour Science</full-title></periodical><alt-periodical><full-title>Appl Anim Behav Sci</full-title></alt-periodical><pages>69-82</pages><volume>96</volume><number>1-2</number><keywords><keyword>prenatal stress</keyword><keyword>chicken</keyword><keyword>fear</keyword><keyword>cognition</keyword><keyword>competition</keyword><keyword>prenatal stress</keyword><keyword>feather pecking</keyword><keyword>plasma-corticosterone</keyword><keyword>pregnancy</keyword><keyword>animals</keyword><keyword>anxiety</keyword><keyword>alters</keyword><keyword>yolk</keyword><keyword>rats</keyword><keyword>hen</keyword></keywords><dates><year>2006</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0168-1591</isbn><accession-num>ISI:000235262600006</accession-num><urls><related-urls><url>&lt;Go to ISI&gt;://000235262600006</url></related-urls></urls><electronic-resource-num>DOI 10.1016/j.applanim.2005.04.020</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_14" \o "Janczak, 2006 #37863" 14]. 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P.</author></authors></contributors><auth-address>Brain and Behaviour Centre, Academia Sinica, Beijing, People&apos;s Republic of China. suinan@ihw.co.cn</auth-address><titles><title>Interactions of corticosterone and embryonic light deprivation on memory retention in day-old chicks</title><secondary-title>Brain Res Dev Brain Res</secondary-title></titles><periodical><full-title>Brain Res Dev Brain Res</full-title></periodical><pages>269-72</pages><volume>101</volume><number>1-2</number><edition>1997/07/18</edition><keywords><keyword>Animals</keyword><keyword>Anti-Inflammatory Agents/*pharmacology</keyword><keyword>Avoidance Learning/drug effects/physiology</keyword><keyword>Chick Embryo</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>*Darkness</keyword><keyword>Memory/*drug effects/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Jul 18</date></pub-dates></dates><isbn>0165-3806 (Print)&#xD;0165-3806 (Linking)</isbn><accession-num>9263600</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9263600</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_19" \o "Sui, 1997 #1722" 19], and increased the rate of pecking behavior at grains and pebbles  ADDIN EN.CITE <EndNote><Cite><Author>Freire</Author><Year>2006</Year><RecNum>37754</RecNum><DisplayText>[20]</DisplayText><record><rec-number>37754</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37754</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Freire, R.</author><author>van Dort, S.</author><author>Rogers, L. J.</author></authors></contributors><auth-address>Centre for Neuroscience and Animal Behaviour, University of New England, Australia. rfreire2@une.edu.au</auth-address><titles><title>Pre- and post-hatching effects of corticosterone treatment on behavior of the domestic chick</title><secondary-title>Horm Behav</secondary-title></titles><periodical><full-title>Horm Behav</full-title></periodical><pages>157-65</pages><volume>49</volume><number>2</number><edition>2005/07/02</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn/*physiology</keyword><keyword>Behavior, Animal/*drug effects</keyword><keyword>Chick Embryo</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>Female</keyword><keyword>Functional Laterality/physiology</keyword><keyword>Hawks</keyword><keyword>Male</keyword><keyword>Photic Stimulation</keyword><keyword>Predatory Behavior</keyword><keyword>Psychomotor Performance/drug effects</keyword></keywords><dates><year>2006</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0018-506X (Print)&#xD;0018-506X (Linking)</isbn><accession-num>15990098</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15990098</url></related-urls></urls><electronic-resource-num>S0018-506X(05)00143-1 [pii]&#xD;10.1016/j.yhbeh.2005.05.015</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_20" \o "Freire, 2006 #37754" 20]. 
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Rensel,Laboratory of Neuroendocrinology, University of California, Los Angeles,United States,mrensel@ucla.edu</auth-address><titles><title>11�-HSD Types 1 and 2 in the Songbird Brain</title><secondary-title>Frontiers in Endocrinology</secondary-title><short-title>11�"HSDs in the Songbird Brain</short-title></titles><periodical><full-title>Frontiers in endocrinology</full-title></periodical><volume>9</volume><number>86</number><keywords><keyword>glucocorticoid,Mineralocorticoid,stress response,songbird,11 beta hydroxysteroid dehydrogenase</keyword></keywords><dates><year>2018</year><pub-dates><date>2018-March-12</date></pub-dates></dates><isbn>1664-2392</isbn><work-type>Original Research</work-type><urls><related-urls><url>https://www.frontiersin.org/article/10.3389/fendo.2018.00086</url></related-urls></urls><electronic-resource-num>10.3389/fendo.2018.00086</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_22" \o "Rensel, 2018 #67240" 22], while 11�-hydroxysteroid dehydrogenase type 2 (11�-HSD2) inactivates GCs  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_23" \o "Diederich, 1998 #8843" 23-27]. In mammals, 11�-HSD1 gene mRNA is expressed mainly in liver  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_28" \o "Zou, 2018 #67245" 28], kidney  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_29" \o "Sagmeister, 2019 #67244" 29] and lung  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_30" \o "Rajan, 1995 #20813" 30,  HYPERLINK \l "_ENREF_31" \o "Chapman, 2013 #14956" 31], whereas, 11HSD2 mostly expressed in kidney  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_32" \o "Sadosky, 2008 #67248" 32], colon  ADDIN EN.CITE <EndNote><Cite><Author>Yang</Author><Year>2013</Year><RecNum>67247</RecNum><DisplayText>[33]</DisplayText><record><rec-number>67247</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67247</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Yang, Shilin</author><author>Jiang, Li</author><author>Zhang, Ming-Zhi</author></authors></contributors><titles><title>11�-Hydroxysteroid Dehydrogenase Type II is a Potential Target for Prevention of Colorectal Tumorigenesis</title><secondary-title>Journal of oncobiomarkers</secondary-title><alt-title>J Oncobiomarkers</alt-title></titles><periodical><full-title>Journal of oncobiomarkers</full-title><abbr-1>J Oncobiomarkers</abbr-1></periodical><alt-periodical><full-title>Journal of oncobiomarkers</full-title><abbr-1>J Oncobiomarkers</abbr-1></alt-periodical><pages>002</pages><volume>1</volume><number>1</number><dates><year>2013</year></dates><isbn>2325-2340</isbn><accession-num>23936870</accession-num><urls><related-urls><url>https://pubmed.ncbi.nlm.nih.gov/23936870</url><url>https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3736594/</url></related-urls></urls><electronic-resource-num>10.13188/2325-2340.1000002</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_33" \o "Yang, 2013 #67247" 33] and placenta  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_34" \o "Albiston, 1994 #21319" 34]. 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Substantial exposure of GCs consider a common relationship between the prenatal surrounding environment  ADDIN EN.CITE <EndNote><Cite><Author>Hartman</Author><Year>2018</Year><RecNum>67266</RecNum><DisplayText>[38]</DisplayText><record><rec-number>67266</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67266</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hartman, Sarah</author><author>Freeman, Sara M.</author><author>Bales, Karen L.</author><author>Belsky, Jay</author></authors></contributors><titles><title>Prenatal Stress as a Risk-and an Opportunity-Factor</title><secondary-title>Psychological science</secondary-title><alt-title>Psychol Sci</alt-title></titles><periodical><full-title>Psychol Sci</full-title><abbr-1>Psychological science</abbr-1></periodical><alt-periodical><full-title>Psychol Sci</full-title><abbr-1>Psychological science</abbr-1></alt-periodical><pages>572-580</pages><volume>29</volume><number>4</number><edition>02/07</edition><keywords><keyword>*anxiety</keyword><keyword>*early environment</keyword><keyword>*plasticity</keyword><keyword>*prenatal stress</keyword><keyword>*vasopressin</keyword><keyword>Amygdala/metabolism</keyword><keyword>Animals</keyword><keyword>Arvicolinae</keyword><keyword>*Behavior, Animal</keyword><keyword>Female</keyword><keyword>Male</keyword><keyword>Oxytocin/*metabolism</keyword><keyword>Pregnancy</keyword><keyword>*Prenatal Exposure Delayed Effects</keyword><keyword>Social Behavior</keyword><keyword>*Stress, Psychological</keyword><keyword>Vasopressins/*metabolism</keyword></keywords><dates><year>2018</year></dates><publisher>SAGE Publications</publisher><isbn>1467-9280&#xD;0956-7976</isbn><accession-num>29412055</accession-num><urls><related-urls><url>https://pubmed.ncbi.nlm.nih.gov/29412055</url><url>https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6350146/</url></related-urls></urls><electronic-resource-num>10.1177/0956797617739983</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_38" \o "Hartman, 2018 #67266" 38], fetal growth  ADDIN EN.CITE <EndNote><Cite><Author>Coussons-Read</Author><Year>2013</Year><RecNum>67251</RecNum><DisplayText>[39]</DisplayText><record><rec-number>67251</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67251</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Coussons-Read, Mary E.</author></authors></contributors><titles><title>Effects of prenatal stress on pregnancy and human development: mechanisms and pathways</title><secondary-title>Obstetric medicine</secondary-title><alt-title>Obstet Med</alt-title></titles><periodical><full-title>Obstetric medicine</full-title><abbr-1>Obstet Med</abbr-1></periodical><alt-periodical><full-title>Obstetric medicine</full-title><abbr-1>Obstet Med</abbr-1></alt-periodical><pages>52-57</pages><volume>6</volume><number>2</number><edition>05/03</edition><keywords><keyword>development</keyword><keyword>inflammation</keyword><keyword>pregnancy</keyword><keyword>stress</keyword></keywords><dates><year>2013</year></dates><publisher>SAGE Publications</publisher><isbn>1753-495X&#xD;1753-4968</isbn><accession-num>27757157</accession-num><urls><related-urls><url>https://pubmed.ncbi.nlm.nih.gov/27757157</url><url>https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5052760/</url></related-urls></urls><electronic-resource-num>10.1177/1753495X12473751</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_39" \o "Coussons-Read, 2013 #67251" 39] and adult neuroendocrine  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_40" \o "Grundwald, 2015 #67252" 40,  HYPERLINK \l "_ENREF_41" \o "Vaeroy, 2019 #67265" 41] and affective disorders  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_42" \o "Welberg, 2000 #15594" 42]. The Inhibition of 11�-HSD1 was reported to prevent stress effects on hippocampal synaptic plasticity  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_43" \o "Sarabdjitsingh, 2014 #15595" 43]and impairs contextual fear behavior  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_43" \o "Sarabdjitsingh, 2014 #15595" 43].
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A.</author></authors></contributors><auth-address>Department of Neurobiology, Harvard Medical School, Boston, MA 02115, USA. edward_kravitz@hms.harvard.edu</auth-address><titles><title>Serotonin and aggression: insights gained from a lobster model system and speculations on the role of amine neurons in a complex behavior</title><secondary-title>J Comp Physiol A</secondary-title></titles><periodical><full-title>J Comp Physiol A</full-title></periodical><pages>221-38</pages><volume>186</volume><number>3</number><edition>2000/04/11</edition><keywords><keyword>Aggression/*physiology</keyword><keyword>Animals</keyword><keyword>*Nephropidae</keyword><keyword>Serotonin/*pharmacology</keyword><keyword>Social Behavior</keyword></keywords><dates><year>2000</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0340-7594 (Print)&#xD;0340-7594 (Linking)</isbn><accession-num>10757238</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10757238</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_45" \o "Kravitz, 2000 #50642" 45]. 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H.</author><author>Winberg, S.</author></authors></contributors><auth-address>Department of Biology, University of South Dakota, Vermillion, SD 57069 USA. cliff@usd.edu</auth-address><titles><title>Interactions between the neural regulation of stress and aggression</title><secondary-title>J Exp Biol</secondary-title></titles><periodical><full-title>J Exp Biol</full-title></periodical><pages>4581-9</pages><volume>209</volume><number>Pt 23</number><edition>2006/11/23</edition><keywords><keyword>Aggression/*physiology</keyword><keyword>Animals</keyword><keyword>Neurons/*physiology</keyword><keyword>Stress, Physiological/*physiopathology</keyword></keywords><dates><year>2006</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>0022-0949 (Print)&#xD;0022-0949 (Linking)</isbn><accession-num>17114393</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=17114393</url></related-urls></urls><electronic-resource-num>209/23/4581 [pii]&#xD;10.1242/jeb.02565</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_46" \o "Summers, 2006 #50610" 46]. 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Low concentrations of blood 5-HT are linked with altered physiological status  ADDIN EN.CITE <EndNote><Cite><Author>Brummelte</Author><Year>2017</Year><RecNum>67260</RecNum><DisplayText>[49]</DisplayText><record><rec-number>67260</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67260</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Brummelte, S.</author><author>Mc Glanaghy, E.</author><author>Bonnin, A.</author><author>Oberlander, T. F.</author></authors></contributors><titles><title>Developmental changes in serotonin signaling: Implications for early brain function, behavior and adaptation</title><secondary-title>Neuroscience</secondary-title><alt-title>Neuroscience</alt-title></titles><periodical><full-title>Neuroscience</full-title><abbr-1>Neuroscience</abbr-1></periodical><alt-periodical><full-title>Neuroscience</full-title><abbr-1>Neuroscience</abbr-1></alt-periodical><pages>212-231</pages><volume>342</volume><edition>02/22</edition><keywords><keyword>*5-HT</keyword><keyword>*SSRI exposure</keyword><keyword>*epigenetics</keyword><keyword>*fetal development</keyword><keyword>*maternal depression</keyword><keyword>*neurodevelopment</keyword><keyword>Animals</keyword><keyword>Brain/*growth &amp; development/*metabolism</keyword><keyword>Humans</keyword><keyword>Serotonin/genetics/*metabolism</keyword><keyword>Signal Transduction/genetics/physiology</keyword></keywords><dates><year>2017</year></dates><isbn>1873-7544&#xD;0306-4522</isbn><accession-num>26905950</accession-num><urls><related-urls><url>https://pubmed.ncbi.nlm.nih.gov/26905950</url><url>https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5310545/</url></related-urls></urls><electronic-resource-num>10.1016/j.neuroscience.2016.02.037</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_49" \o "Brummelte, 2017 #67260" 49], including the HPA axis  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_50" \o "Leonard, 2006 #49398" 50] and aggressive behavior in humans  ADDIN EN.CITE <EndNote><Cite><Author>Booij</Author><Year>2010</Year><RecNum>49402</RecNum><DisplayText>[51]</DisplayText><record><rec-number>49402</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">49402</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Booij, L.</author><author>Tremblay, R. 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L.</author><author>Durand, F.</author><author>Diksic, M.</author><author>Turecki, G.</author><author>Benkelfat, C.</author></authors></contributors><auth-address>Department of Psychiatry, McGill University, Montreal, Canada.</auth-address><titles><title>Brain serotonin synthesis in adult males characterized by physical aggression during childhood: a 21-year longitudinal study</title><secondary-title>PLoS ONE</secondary-title></titles><periodical><full-title>PLoS ONE</full-title></periodical><pages>e11255</pages><volume>5</volume><number>6</number><edition>2010/06/29</edition><keywords><keyword>Adult</keyword><keyword>*Aggression</keyword><keyword>Brain/*metabolism/radionuclide imaging</keyword><keyword>Emotions</keyword><keyword>Humans</keyword><keyword>Longitudinal Studies</keyword><keyword>Male</keyword><keyword>Memory</keyword><keyword>Positron-Emission Tomography</keyword><keyword>Serotonin/*biosynthesis</keyword><keyword>Tryptophan/blood</keyword></keywords><dates><year>2010</year></dates><isbn>1932-6203 (Electronic)&#xD;1932-6203 (Linking)</isbn><accession-num>20582306</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=20582306</url></related-urls></urls><custom2>2889822</custom2><electronic-resource-num>10.1371/journal.pone.0011255</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_51" \o "Booij, 2010 #49402" 51]. In humans, childhood stress is found to induce aggression which associated with disrupted HPA axis function  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_52" \o "da Cunha-Bang, 2016 #67264" 52] and reduced the 5-HT system function in adulthood  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_53" \o "Veenema, 2009 #49441" 53,  HYPERLINK \l "_ENREF_54" \o "Shah, 2018 #67263" 54]. The frequency of aggressive behaviors increased in the hen of �Dekalb XL (DXL) and low group productivity and survivability (LGPS) hen treated with 5-hydroxytryptamine (serotonin) receptor 1A (5-HT1A) antagonist indicating that serotonin plays a major role in aggressive behaviors  ADDIN EN.CITE <EndNote><Cite><Author>K�stner</Author><Year>2019</Year><RecNum>67268</RecNum><DisplayText>[55]</DisplayText><record><rec-number>67268</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67268</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>K�stner, Niklas</author><author>Richter, S. 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    The mode of action of GCs in the cells is mediated through glucocorticoid receptors (GR), and mineralocorticoid receptors (MR)  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_58" \o "Tsugita, 2009 #6813" 58]. The intracellular availability of active GC is modulated by pre-receptor mechanisms  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_21" \o "Edwards, 1996 #9766" 21] and corticosterone binding globulin (CBG). The 11�-hydroxysteroid dehydrogenase (11�-HSD1) activates, whereas 11�-hydroxysteroid dehydrogenase (11�-HSD2) deactivates GCs in all animal species  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_23" \o "Diederich, 1998 #8843" 23-27]. In addition, in birds, 20-hydroxysteroid dehydrogenase (20-HSD) is an abundantly and ubiquitously expressed enzyme  ADDIN EN.CITE <EndNote><Cite><Author>Rensel</Author><Year>2018</Year><RecNum>67240</RecNum><DisplayText>[22]</DisplayText><record><rec-number>67240</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67240</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rensel,Michelle A.</author><author>Ding,Jessica A.</author><author>Pradhan,Devaleena S.</author><author>Schlinger,Barney A.</author></authors></contributors><auth-address>Michelle A. Rensel,The Institute for Society and Genetics, University of California, Los Angeles,United States,mrensel@ucla.edu&#xD;Michelle A. Rensel,Laboratory of Neuroendocrinology, University of California, Los Angeles,United States,mrensel@ucla.edu</auth-address><titles><title>11�-HSD Types 1 and 2 in the Songbird Brain</title><secondary-title>Frontiers in Endocrinology</secondary-title><short-title>11�"HSDs in the Songbird Brain</short-title></titles><periodical><full-title>Frontiers in endocrinology</full-title></periodical><volume>9</volume><number>86</number><keywords><keyword>glucocorticoid,Mineralocorticoid,stress response,songbird,11 beta hydroxysteroid dehydrogenase</keyword></keywords><dates><year>2018</year><pub-dates><date>2018-March-12</date></pub-dates></dates><isbn>1664-2392</isbn><work-type>Original Research</work-type><urls><related-urls><url>https://www.frontiersin.org/article/10.3389/fendo.2018.00086</url></related-urls></urls><electronic-resource-num>10.3389/fendo.2018.00086</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_22" \o "Rensel, 2018 #67240" 22], which transforms GCs to the inactive 20-dihydrocorticosterone  ADDIN EN.CITE <EndNote><Cite><Author>Kucka</Author><Year>2006</Year><RecNum>1292</RecNum><DisplayText>[59]</DisplayText><record><rec-number>1292</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">1292</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kucka, M.</author><author>Vagnerova, K.</author><author>Klusonova, P.</author><author>Miksik, I.</author><author>Pacha, J.</author></authors></contributors><auth-address>Institute of Physiology, Czech Academy of Sciences, Videnska 1083, CZ-142 20 Prague 4, Czech Republic.</auth-address><titles><title>Corticosterone metabolism in chicken tissues: evidence for tissue-specific distribution of steroid dehydrogenases</title><secondary-title>Gen Comp Endocrinol</secondary-title></titles><periodical><full-title>Gen Comp Endocrinol</full-title></periodical><pages>377-83</pages><volume>147</volume><number>3</number><edition>2006/04/01</edition><keywords><keyword>11-beta-Hydroxysteroid Dehydrogenases/*metabolism</keyword><keyword>20-Hydroxysteroid Dehydrogenases/*metabolism</keyword><keyword>Animals</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*metabolism</keyword><keyword>Organ Specificity</keyword><keyword>*Tissue Distribution</keyword></keywords><dates><year>2006</year><pub-dates><date>Jul</date></pub-dates></dates><isbn>0016-6480 (Print)&#xD;0016-6480 (Linking)</isbn><accession-num>16574115</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=16574115</url></related-urls></urls><electronic-resource-num>S0016-6480(06)00046-3 [pii]&#xD;10.1016/j.ygcen.2006.02.007</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_59" \o "Kucka, 2006 #1292" 59]. GCs are reported to increase aggressive behavior through both genomic  ADDIN EN.CITE <EndNote><Cite><Author>Veenit</Author><Year>2013</Year><RecNum>67271</RecNum><DisplayText>[60]</DisplayText><record><rec-number>67271</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67271</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Veenit,Vandana</author><author>Cordero,Mar�a</author><author>Tzanoulinou,Stamatina</author><author>Sandi,Carmen</author></authors></contributors><auth-address>Prof Carmen Sandi,Ecole Polytechnique Federal Lausanne,Lausanne,Switzerland,carmen.sandi@epfl.ch</auth-address><titles><title>Increased corticosterone in peripubertal rats leads to long-lasting alterations in social exploration and aggression</title><secondary-title>Frontiers in Behavioral Neuroscience</secondary-title><short-title>Peripubertal corticosterone alters social behaviors</short-title></titles><periodical><full-title>Frontiers in behavioral neuroscience</full-title><abbr-1>Front Behav Neurosci</abbr-1></periodical><volume>7</volume><number>26</number><keywords><keyword>Corticosterone,Puberty,adolescence,stress,sociability,aggressive behavior,emotion</keyword></keywords><dates><year>2013</year><pub-dates><date>2013-April-04</date></pub-dates></dates><isbn>1662-5153</isbn><work-type>Original Research</work-type><urls><related-urls><url>https://www.frontiersin.org/article/10.3389/fnbeh.2013.00026</url></related-urls></urls><electronic-resource-num>10.3389/fnbeh.2013.00026</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_60" \o "Veenit, 2013 #67271" 60] and non genomic mechanisms  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_61" \o "Mikics, 2004 #15568" 61-63]. The action of GCs is controlled at the hypothalamic and pituitary level  ADDIN EN.CITE <EndNote><Cite><Author>Smith</Author><Year>2006</Year><RecNum>8543</RecNum><DisplayText>[64]</DisplayText><record><rec-number>8543</rec-number><foreign-keys><key app="EN" db-id="eww2eazad2dr9neetrmxespbeste2wde2xr0">8543</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Smith, S. M.</author><author>Vale, W. W.</author></authors></contributors><auth-address>Clayton Foundation Laboratories for Peptide Biology, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA.</auth-address><titles><title>The role of the hypothalamic-pituitary-adrenal axis in neuroendocrine responses to stress</title><secondary-title>Dialogues Clin Neurosci</secondary-title></titles><periodical><full-title>Dialogues Clin Neurosci</full-title></periodical><pages>383-95</pages><volume>8</volume><number>4</number><edition>2007/02/13</edition><keywords><keyword>Animals</keyword><keyword>Endocrine Glands/physiopathology</keyword><keyword>Humans</keyword><keyword>Hypothalamo-Hypophyseal System/pathology/*physiopathology</keyword><keyword>Limbic System/physiopathology</keyword><keyword>Neurons/physiology</keyword><keyword>Neurosecretory Systems/*physiopathology</keyword><keyword>Pituitary-Adrenal System/pathology/*physiopathology</keyword><keyword>Stress, Physiological/*physiopathology</keyword><keyword>Sympathetic Nervous System/physiopathology</keyword></keywords><dates><year>2006</year></dates><isbn>1294-8322 (Print)&#xD;1294-8322 (Linking)</isbn><accession-num>17290797</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=17290797</url></related-urls></urls><custom2>3181830</custom2><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_64" \o "Smith, 2006 #8543" 64]. One important regulatory mechanism consists of modulation in the expression of the two isoforms of 11�-HSDs which catalyze the inter-conversion of GCs  ADDIN EN.CITE <EndNote><Cite><Author>Jun</Author><Year>2013</Year><RecNum>15587</RecNum><DisplayText>[65]</DisplayText><record><rec-number>15587</rec-number><foreign-keys><key app="EN" db-id="eww2eazad2dr9neetrmxespbeste2wde2xr0">15587</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Jun, Young Joon</author><author>Park, Se Jin</author><author>Kim, Tae Hoon</author><author>Lee, Seung Hoon</author><author>Lee, Ki Jeong</author><author>Hwang, Soo Min</author><author>Lee, Sang Hag</author></authors></contributors><titles><title>Expression of 11�-hydroxysteroid dehydrogenase 1�and�2�in�patients with chronic rhinosinusitis and their possible contribution to local glucocorticoid activation in�sinus mucosa</title><secondary-title>Journal of Allergy and Clinical Immunology</secondary-title></titles><periodical><full-title>Journal of Allergy and Clinical Immunology</full-title></periodical><pages>926-934.e6</pages><volume>134</volume><number>4</number><dates><year>2013</year></dates><publisher>Elsevier</publisher><work-type>doi: 10.1016/j.jaci.2014.03.033</work-type><urls><related-urls><url>http://www.jacionline.org/article/S0091-6749(14)00514-4/abstract</url></related-urls></urls><electronic-resource-num>10.1016/j.jaci.2014.03.033</electronic-resource-num><access-date>2014/11/21</access-date></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_65" \o "Jun, 2013 #15587" 65]. Several reports have indicated that the hypothalamus has a role in aggression in different species inducing finches  ADDIN EN.CITE <EndNote><Cite><Author>Goodson</Author><Year>2012</Year><RecNum>15543</RecNum><DisplayText>[66]</DisplayText><record><rec-number>15543</rec-number><foreign-keys><key app="EN" db-id="eww2eazad2dr9neetrmxespbeste2wde2xr0">15543</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Goodson, J. L.</author><author>Kelly, A. M.</author><author>Kingsbury, M. A.</author><author>Thompson, R. R.</author></authors></contributors><auth-address>Department of Biology, Indiana University, Bloomington IN 47405, USA. jlgoodso@indiana.edu</auth-address><titles><title>An aggression-specific cell type in the anterior hypothalamus of finches</title><secondary-title>Proc Natl Acad Sci U S A</secondary-title></titles><periodical><full-title>Proc Natl Acad Sci U S A</full-title></periodical><pages>13847-52</pages><volume>109</volume><number>34</number><edition>2012/08/09</edition><keywords><keyword>Aggression/*physiology</keyword><keyword>Animals</keyword><keyword>Behavior, Animal/physiology</keyword><keyword>Brain Mapping/*methods</keyword><keyword>Female</keyword><keyword>Finches/*physiology</keyword><keyword>Gene Expression Regulation</keyword><keyword>Hypothalamus/metabolism/*physiology</keyword><keyword>Male</keyword><keyword>Models, Biological</keyword><keyword>Neurons/metabolism</keyword><keyword>Neuropeptides/chemistry</keyword><keyword>Songbirds/metabolism/physiology</keyword><keyword>Territoriality</keyword><keyword>Vocalization, Animal/physiology</keyword></keywords><dates><year>2012</year><pub-dates><date>Aug 21</date></pub-dates></dates><isbn>1091-6490 (Electronic)&#xD;0027-8424 (Linking)</isbn><accession-num>22872869</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=22872869</url></related-urls></urls><custom2>3427066</custom2><electronic-resource-num>1207995109 [pii]&#xD;10.1073/pnas.1207995109</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_66" \o "Goodson, 2012 #15543" 66], sparrows  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_67" \o "Mukai, 2009 #15547" 67] and rats  ADDIN EN.CITE <EndNote><Cite><Author>Kruk</Author><Year>1998</Year><RecNum>15549</RecNum><DisplayText>[68]</DisplayText><record><rec-number>15549</rec-number><foreign-keys><key app="EN" db-id="eww2eazad2dr9neetrmxespbeste2wde2xr0">15549</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kruk, M. R.</author><author>Westphal, K. G.</author><author>Van Erp, A. M.</author><author>van Asperen, J.</author><author>Cave, B. J.</author><author>Slater, E.</author><author>de Koning, J.</author><author>Haller, J.</author></authors></contributors><auth-address>Medical Pharmacology, Leiden-Amsterdam Center for Drug Research, The Netherlands.</auth-address><titles><title>The hypothalamus: cross-roads of endocrine and behavioural regulation in grooming and aggression</title><secondary-title>Neurosci Biobehav Rev</secondary-title></titles><periodical><full-title>Neurosci Biobehav Rev</full-title></periodical><pages>163-77</pages><volume>23</volume><number>2</number><edition>1999/01/12</edition><keywords><keyword>Aggression/*physiology/*psychology</keyword><keyword>Animals</keyword><keyword>Behavior, Animal/*physiology</keyword><keyword>Endocrine Glands/*physiology</keyword><keyword>Grooming/*physiology</keyword><keyword>Hypothalamus/*physiology</keyword></keywords><dates><year>1998</year></dates><isbn>0149-7634 (Print)&#xD;0149-7634 (Linking)</isbn><accession-num>9884110</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9884110</url></related-urls></urls><electronic-resource-num>S0149763498000189 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_68" \o "Kruk, 1998 #15549" 68]. Yet, the effects of embryonic CORT exposure on hypothalamic glucocrticoid metabolic enzyme gene, 5-HT receptor expression and its association with aggressive behavior in the of chicken is not reported. 
    In the present study we used a model of in ovo injection of CORT before incubation to test our hypotheses that aggressive behavior and plasma CORT concentration may be influenced by CORT treatment and these changes may be associated with hypothalamic GCs metabolic enzymes gene and 5-HT receptor expression. 

Materials and Methods
Egg incubation and CORT injection

    Two hundred and ten fertilized chicken eggs overall mean mass (64.6 � 0.44 g) were selected from eggs laid by hens one month after onset of lay and randomly divided into three groups (70 in each group). CORT (Sigma-Aldrich, USA) was dissolved in absolute alcohol, rather than the oil that affected embryonic development of chickens in our previous trials, and diluted in PBS to produce doses of 0.2 (g and 1 (g in a volume of 100 �L solution containing a minimal amount of alcohol. The high and low CORT dose was determined based on previous publications  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_69" \o "Heiblum, 2001 #870" 69,  HYPERLINK \l "_ENREF_70" \o "Haussmann, 2012 #34578" 70]; taking into consideration the CORT concentration detected in the yolk (3-4 ng/g) and the albumen (0.5 ng/g)  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_71" \o "Ahmed, 2013 #6760" 71]. Before incubation, the eggs were injected with PBS (control) and a 0.2 (g (low) or a 1 (g (high) dose of CORT under aseptic conditions. Eggs were injected randomly by advancing a Hamilton syringe into a hole in the middle of the long axis until the yolk membrane was penetrated (approximately 20 mm below the surface). The incubation conditions were set according to our previous publication  ADDIN EN.CITE <EndNote><Cite><Author>Su</Author><Year>2012</Year><RecNum>37419</RecNum><DisplayText>[72]</DisplayText><record><rec-number>37419</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37419</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Su, L.</author><author>Rao, K.</author><author>Guo, F.</author><author>Li, X.</author><author>Ahmed, A. A.</author><author>Ni, Y.</author><author>Grossmann, R.</author><author>Zhao, R.</author></authors></contributors><auth-address>Key Laboratory of Animal Physiology and Biochemistry, Ministry of Agriculture, Nanjing Agricultural University, Nanjing 210095, China; College of Bioengineering, Henan University of Technology, Zhengzhou, 450001, China.</auth-address><titles><title>In ovo leptin administration inhibits chorioallantoic membrane angiogenesis in female chicken embryos through the STAT3-mediated vascular endothelial growth factor (VEGF) pathway</title><secondary-title>Domest Anim Endocrinol</secondary-title></titles><periodical><full-title>Domest Anim Endocrinol</full-title></periodical><edition>2012/03/16</edition><dates><year>2012</year><pub-dates><date>Mar 3</date></pub-dates></dates><isbn>1879-0054 (Electronic)&#xD;0739-7240 (Linking)</isbn><accession-num>22417645</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=22417645</url></related-urls></urls><electronic-resource-num>S0739-7240(12)00009-4 [pii]&#xD;10.1016/j.domaniend.2012.01.007</electronic-resource-num><language>Eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_72" \o "Su, 2012 #37419" 72]. Chicks were hatched inside the incubator and were left to dry completely (up to 12 h) before they were removed. The hatchability of the eggs ranged from70% to 75% and no obvious differences in hatchability or hatching time were observed among three groups. One-day-old chicks were individually weighed, wing banded, and placed into battery cages with continuous fluorescent lighting. The temperature was adjusted to 32�35�%C during the first week, and reduced approximately 3�%C per week until 21�%C. Both sexes were transferred to floor pens covered with sawdust litter. The stock density was 20 25 kg/m2. The relative humidity was maintained at 40 60%, and the lighting, ventilation, as well as the feeding and management procedures complied with the Feeding Management Regulations of Yellow-feathered Chicken (NY/T 1871-2010). The growth performance was started on Day 1 (D1) posthatch and recorded weekly from hatching to 10 weeks of age. On D133, hens started to lay eggs. On D175, blood samples were collected for plasma CORT measurement. Behaviors test were performed on posthatch D196. On D210 tonic immobility tests were performed twice per day for three days using different batches of chickens. We used 6 animals per group, in total 18 animals for parameters except the growth rate. We tried not to use the same batch of 6 birds for different measurements in order to minimize the stress caused by different manipulations. On D245, all chickens were killed by rapid decapitation one of the physical methods, which have been used as an ethical type of euthanasia. The hypothalami were collected, washed with PBS then put in liquid nitrogen and later kept at -80 �C for further analysis. The experiment procedures were approved by the Animal Ethics Committee of Nanjing Agricultural University
   
Aggressive behavior test
    The behavior tests were performed on D196 as described previously  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_73" \o "Kitaysky, 2003 #31" 73]. Briefly, 30 chickens from each group which were unfamiliar to each other from different brooders were placed in an experimental arena (similar in size and structure to their brooders where chickens have been raised) which was established in a room familiar to the animals. The room was visually and acoustically isolated from the aviary. For visual identification, chickens were marked with different colors (red, green, blue) on different locations (head, back and tail). Neither the colors nor their locations affected the behaviors of chickens in the present study. The chicken's behavior was videotaped during a 60 min period. The number of aggressive attacks of each individual was recorded, and aggression was defined as a chicken pecking, grabbing, twisting skin on the head and nape of the other chicken. The observer who recorded and analyzed the aggressive behaviour was not aware of the experimental treatments.

Tonic immobility (TI) test
    TI tests were measured on D210 using different chickens. The TI tests were measured according to the method described previously  ADDIN EN.CITE <EndNote><Cite><Author>Mills</Author><Year>1991</Year><RecNum>48374</RecNum><DisplayText>[74]</DisplayText><record><rec-number>48374</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">48374</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Mills, A. D.</author><author>Faure, J. M.</author></authors></contributors><auth-address>Station de Recherches Avicoles, Institut National de la Recherche Agronomique, Centre de Tours-Nouzilly, Nouzilly, France.</auth-address><titles><title>Divergent selection for duration of tonic immobility and social reinstatement behavior in Japanese quail (Coturnix coturnix japonica) chicks</title><secondary-title>J Comp Psychol</secondary-title></titles><periodical><full-title>J Comp Psychol</full-title></periodical><pages>25-38</pages><volume>105</volume><number>1</number><edition>1991/03/01</edition><keywords><keyword>Animals</keyword><keyword>Arousal/*genetics</keyword><keyword>Coturnix/*genetics</keyword><keyword>*Fear</keyword><keyword>Habituation, Psychophysiologic/genetics</keyword><keyword>Motor Activity/genetics</keyword><keyword>*Selection, Genetic</keyword><keyword>*Social Environment</keyword></keywords><dates><year>1991</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0735-7036 (Print)&#xD;0021-9940 (Linking)</isbn><accession-num>2032452</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=2032452</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_74" \o "Mills, 1991 #48374" 74]. Briefly, a chicken was carried individually to another isolated room devoid of other birds. The chicken was placed on its back on the floor and restrained for at least 20 s (with one hand on the sternum and one lightly cupping the head of the bird). The experimenter remained silent and virtually motionless in the room, out of the bird's sight. The TI duration was considered between 10 and 600 s. If the chicken terminated in <10 s, it was captured, and the trial was repeated. If TI was not attained after 3 attempts, a score of 0 s was given. Conversely, if the bird failed to right itself after 10 min, the test was terminated and a maximum score of 600 s was given for tonic immobility duration. 
Plasma CORT assay
    The birds used for taking blood samples were trained prior to the sampling to get used to human manipulations. Approximately 1 mL of blood was collected from the jugular vein and duplicate plasma samples (2 x 50 �L) were used for the CORT assay. The plasma CORT concentration was measured with a commercial enzyme immunoassay kit (500655, Cayman Chemical Company, Ann Arbor, MI, USA) according to the manufacturer�s instructions. The calculated detection limit of the assay was 27 pg/mL and all the determinations fell within the range of detection. The intra-assay coefficient of variation was 5%. The cross-reactivity of the antibody was 11% with 11-dehydrocorticosterone, 7% with 11-deoxycorticosterone, 0.31% with progesterone, 0.17% with cortisol, 0.06% with aldosterone, 0.03% with testosterone, 0.02% with pregnenolone, 0.01% with 5�-DHT and less than 0.01% with other steroids. 

RNA extraction and mRNA quantification with Real-time PCR
    Hypothalamus samples were ground with pestle and mortar in liquid N2 and a portion of approximately 100 mg was used for the RNA extraction using the TRIzol total RNA kit (Invitrogen, Biotechnology Co, Ltd, Carlsbad, CA, USA) according to the manufacturer�s instructions, and reverse transcript to cDNA using 0.5 �g/�L (4�L contains 4�g) of RNA with the PrimeScript RT reagent kit according to the manufacturers instruction (Takara). To investigate the effect of the in ovo injection of CORT on the expression of hypothalamic genes, real-time PCR was performed in an Mx3000P (Stratagene, USA) according to published methods  ADDIN EN.CITE <EndNote><Cite><Author>Li</Author><Year>2011</Year><RecNum>19876</RecNum><DisplayText>[75]</DisplayText><record><rec-number>19876</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">19876</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Li, R.</author><author>Hu, Y.</author><author>Ni, Y.</author><author>Xia, D.</author><author>Grossmann, R.</author><author>Zhao, R.</author></authors></contributors><auth-address>Key Laboratory of Animal Physiology and Biochemistry, Nanjing Agricultural University, Nanjing 210095, China.</auth-address><titles><title>Leptin stimulates hepatic activation of thyroid hormones and promotes early posthatch growth in the chicken</title><secondary-title>Comp Biochem Physiol A Mol Integr Physiol</secondary-title></titles><periodical><full-title>Comp Biochem Physiol A Mol Integr Physiol</full-title></periodical><pages>200-6</pages><volume>160</volume><number>2</number><edition>2011/06/18</edition><dates><year>2011</year><pub-dates><date>Oct</date></pub-dates></dates><isbn>1531-4332 (Electronic)&#xD;1095-6433 (Linking)</isbn><accession-num>21679771</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=21679771</url></related-urls></urls><electronic-resource-num>S1095-6433(11)00169-3 [pii]&#xD;10.1016/j.cbpa.2011.06.001</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_75" \o "Li, 2011 #19876" 75]. Mock RT and No Template Controls (NTC) were included to monitor the possible contamination of genomic and environmental DNA at the RT and PCR steps. A pooled sample made by mixing equal quantities of the RT products (cDNA) from all the samples was used for optimizing the PCR conditions and tailoring the standard curves for each target gene, and melting curves were performed to insure a single specific PCR product for each gene. The PCR products were sequenced to validate the identity of the amplicons. Primers specific for the 11�-HSD1, 11�-HSD2, 20-HSD, GR, MR, 5-HTR1A and 5-HTR1B (Table 2) were synthesized by Geneary, Shanghai, China. Chicken �-actin was used as a reference gene for normalization purposes. The method of 2"��Ct was used to analyze the real-time PCR data  ADDIN EN.CITE <EndNote><Cite><Author>Livak</Author><Year>2001</Year><RecNum>25138</RecNum><DisplayText>[76]</DisplayText><record><rec-number>25138</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">25138</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Livak, K. J.</author><author>Schmittgen, T. D.</author></authors></contributors><auth-address>Applied Biosystems, Foster City, California 94404, USA.</auth-address><titles><title>Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method</title><secondary-title>Methods</secondary-title></titles><periodical><full-title>Methods</full-title></periodical><pages>402-8</pages><volume>25</volume><number>4</number><edition>2002/02/16</edition><keywords><keyword>Algorithms</keyword><keyword>Brain/metabolism</keyword><keyword>Cell Line</keyword><keyword>DNA, Complementary/metabolism</keyword><keyword>Humans</keyword><keyword>Polymerase Chain Reaction/*methods</keyword><keyword>*Reverse Transcriptase Polymerase Chain Reaction</keyword><keyword>Time Factors</keyword></keywords><dates><year>2001</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1046-2023 (Print)&#xD;1046-2023 (Linking)</isbn><accession-num>11846609</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=11846609</url></related-urls></urls><electronic-resource-num>10.1006/meth.2001.1262&#xD;S1046-2023(01)91262-9 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_76" \o "Livak, 2001 #25138" 76].

Statistical analysis 
    Descriptive statistics was performed to check the normality and homogeneity of variances before using parametric analyses. The behavioral data were not normally distributed, so Log 10 transformation was performed before statistical analysis. Body weight was analyzed by repeated measures ANOVA in the General Linear Model (GLM) procedure of SPSS 16.0 (SPSS Inc., Chicago, IL, USA). Behavioral and plasma CORT, as well as the relative quantitative data of gene expression were analyzed by one-way ANOVA using SPSS 16.0 for Windows, followed by a least-significant difference (LSD) test for individual comparisons. A P-value d"0.05 was considered significant.


Results
Growth Rate 
    In ovo injection of CORT significantly (P<0.05) affected the posthatch growth rate of the chickens. The chickens exposed to high doses of CORT grew slower compared to those in the low dose and control groups. As a result, high CORT birds did not weigh as much by the age of 7 weeks (Fig. 1A). 

Aggressive behavior test and tonic immobility test
    CORT administration significantly (P<0.05) increased the frequency of pecking behavior compared to control. The high dose of CORT significantly increased pecking behavior compared to the low group. No significant differences were observed in grabbing and twisting behaviors (Fig. 1B). CORT administration significantly (P<0.05) increased TI duration both in low and high doses of CORT compared to the control group (Fig. 1C). No significant differences were found between the male and the female chickens in the frequency of aggressive behaviors or tonic immobility (data not shown).

CORT Concentrations in plasma
    The high dose of CORT significantly (P<0.05) increased plasma CORT concentration compared to the low dose and control groups. However, the low dose of CORT treatment did not change plasma CORT concentrations (Fig. 1D).

Hypothalamic 11�-HSD1, 11�-HSD2, 20-HSD, GR and MR mRNA expression
    The high dose of CORT treatment significantly increased (P<0.05) hypothalamic 11�-HSD1 mRNA expression (Fig. 2A) whereas, it decreased 11�-HSD2 mRNA (Fig. 2B). In addition, both CORT treatments significantly (P<0.05) decreased MR mRNA expression in the hypothalamus compared to control (Fig. 2E). CORT in ovo did not change neither 20-HSD mRNA (Fig. 2C) nor GR (Fig. 2D) expression in the hypothalamus. 

Hypothalamic 5-HTR1A and 5-HTR1B mRNA expression
    The high dose CORT treatment significantly (P<0.05) increased the hypothalamic expression of 5-HTR1A mRNA compared to the low dose and control groups (Fig. 3A). However, CORT treatment did not affect the 5-HTR1B mRNA expression in the hypothalamus (Fig. 3B).


Discussion
    Ecological factors, such as maternal stress during�the embryonic development increases the aggressive behavior of their offspring in rats and humans  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_77" \o "Monk, 2012 #15589" 77,  HYPERLINK \l "_ENREF_78" \o "Champagne, 2006 #15588" 78]. In avian species, egg CORT elevation was found to increased aggression in captive black-legged kittiwake� ADDIN EN.CITE <EndNote><Cite><Author>Kitaysky</Author><Year>2003</Year><RecNum>31</RecNum><DisplayText>[73]</DisplayText><record><rec-number>31</rec-number><foreign-keys><key app="EN" db-id="s95rxpts7ftxzeetsrn5ad2ftvxep0ax0avf">31</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kitaysky, A. S.</author><author>Kitaiskaia, E.</author><author>Piatt, J.</author><author>Wingfield, J. C.</author></authors></contributors><auth-address>Kitaysky, AS&#xD;Univ Washington, Dept Zool, 24 Kincaid Hall,Box 351800, Seattle, WA 98195 USA&#xD;Univ Washington, Dept Zool, 24 Kincaid Hall,Box 351800, Seattle, WA 98195 USA&#xD;Univ Washington, Dept Zool, Seattle, WA 98195 USA&#xD;US Geol Survey, Alaska Biol Serv Ctr, Anchorage, AK 99503 USA</auth-address><titles><title>Benefits and costs of increased levels of corticosterone in seabird chicks</title><secondary-title>Hormones and Behavior</secondary-title><alt-title>Horm Behav</alt-title></titles><pages>140-149</pages><volume>43</volume><number>1</number><keywords><keyword>aggression</keyword><keyword>associative learning</keyword><keyword>begging</keyword><keyword>cognition</keyword><keyword>corticosterone</keyword><keyword>environmental stress</keyword><keyword>kittiwakes</keyword><keyword>juvenile life-history traits</keyword><keyword>legged kittiwake chicks</keyword><keyword>blue-footed booby</keyword><keyword>stress-response</keyword><keyword>resource-allocation</keyword><keyword>sibling aggression</keyword><keyword>hypothesis</keyword><keyword>asymmetry</keyword><keyword>hunger</keyword><keyword>growth</keyword></keywords><dates><year>2003</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0018-506X</isbn><accession-num>ISI:000182658400018</accession-num><urls><related-urls><url>&lt;Go to ISI&gt;://000182658400018</url></related-urls></urls><electronic-resource-num>Doi 10.1016/S0018-506x(02)00030-2</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_73" \o "Kitaysky, 2003 #31" 73] and increased pecking activities in domestic fowl  ADDIN EN.CITE <EndNote><Cite><Author>Freire</Author><Year>2006</Year><RecNum>15</RecNum><DisplayText>[20]</DisplayText><record><rec-number>15</rec-number><foreign-keys><key app="EN" db-id="s95rxpts7ftxzeetsrn5ad2ftvxep0ax0avf">15</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Freire, R.</author><author>van Dort, S.</author><author>Rogers, L. J.</author></authors></contributors><auth-address>Centre for Neuroscience and Animal Behaviour, University of New England, Australia. rfreire2@une.edu.au</auth-address><titles><title>Pre- and post-hatching effects of corticosterone treatment on behavior of the domestic chick</title><secondary-title>Horm Behav</secondary-title></titles><pages>157-65</pages><volume>49</volume><number>2</number><edition>2005/07/02</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn/*physiology</keyword><keyword>Behavior, Animal/*drug effects</keyword><keyword>Chick Embryo</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>Female</keyword><keyword>Functional Laterality/physiology</keyword><keyword>Hawks</keyword><keyword>Male</keyword><keyword>Photic Stimulation</keyword><keyword>Predatory Behavior</keyword><keyword>Psychomotor Performance/drug effects</keyword></keywords><dates><year>2006</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0018-506X (Print)&#xD;0018-506X (Linking)</isbn><accession-num>15990098</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15990098</url></related-urls></urls><electronic-resource-num>S0018-506X(05)00143-1 [pii]&#xD;10.1016/j.yhbeh.2005.05.015</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_20" \o "Freire, 2006 #37754" 20]. In the present study, the in ovo injection of high dose of CORT significantly decreased growth rate in post-hatch chicks associated with plasma CORT concentrations. The increases of plasma CORT levels this study could be linked and be the cause of growth retardation. These findings in line with previous reports that embryonic CORT exposure retarded the growth in chickens  ADDIN EN.CITE <EndNote><Cite><Author>Heiblum</Author><Year>2001</Year><RecNum>870</RecNum><DisplayText>[69]</DisplayText><record><rec-number>870</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">870</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Heiblum, R.</author><author>Arnon, E.</author><author>Chazan, G.</author><author>Robinzon, B.</author><author>Gvaryahu, G.</author><author>Snapir, N.</author></authors></contributors><auth-address>Department of Animal Sciences, Faculty of Agricultural, The Hebrew University of Jerusalem, Rehovot, Israel. heiblum@agri.huji.ac.il</auth-address><titles><title>Glucocorticoid administration during incubation: embryo mortality and posthatch growth in chickens</title><secondary-title>Poult Sci</secondary-title></titles><periodical><full-title>Poult Sci</full-title></periodical><pages>1357-63</pages><volume>80</volume><number>9</number><edition>2001/09/18</edition><keywords><keyword>Animals</keyword><keyword>Body Weight</keyword><keyword>Chick Embryo/*drug effects/growth &amp; development</keyword><keyword>Corticosterone/administration &amp; dosage/*pharmacology</keyword><keyword>Dose-Response Relationship, Drug</keyword><keyword>Glucocorticoids</keyword><keyword>Hydrocortisone/administration &amp; dosage/*analogs &amp;</keyword><keyword>derivatives/*pharmacology</keyword><keyword>Incubators</keyword><keyword>Lethal Dose 50</keyword><keyword>Time Factors</keyword></keywords><dates><year>2001</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0032-5791 (Print)&#xD;0032-5791 (Linking)</isbn><accession-num>11558923</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=11558923</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_69" \o "Heiblum, 2001 #870" 69] and quail  ADDIN EN.CITE <EndNote><Cite><Author>Hayward</Author><Year>2004</Year><RecNum>1602</RecNum><DisplayText>[8]</DisplayText><record><rec-number>1602</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">1602</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hayward, L. S.</author><author>Wingfield, J. C.</author></authors></contributors><auth-address>Department of Zoology, University of Washington, Seattle, WA 98195, USA. lhayward@u.washington.edu</auth-address><titles><title>Maternal corticosterone is transferred to avian yolk and may alter offspring growth and adult phenotype</title><secondary-title>Gen Comp Endocrinol</secondary-title></titles><periodical><full-title>Gen Comp Endocrinol</full-title></periodical><pages>365-71</pages><volume>135</volume><number>3</number><edition>2004/01/16</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn</keyword><keyword>Birth Weight/drug effects</keyword><keyword>Corticosterone/blood/metabolism/*pharmacology</keyword><keyword>Coturnix/growth &amp; development/*metabolism</keyword><keyword>Egg Yolk/chemistry/*metabolism</keyword><keyword>Female</keyword><keyword>Male</keyword><keyword>Reproduction/drug effects</keyword><keyword>Stress, Physiological/metabolism</keyword><keyword>Zygote/chemistry/metabolism</keyword></keywords><dates><year>2004</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0016-6480 (Print)&#xD;0016-6480 (Linking)</isbn><accession-num>14723888</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=14723888</url></related-urls></urls><electronic-resource-num>S0016648003004039 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_8" \o "Hayward, 2004 #1602" 8], and increased plasma CORT concentration in the chickens  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_14" \o "Janczak, 2006 #37863" 14,  HYPERLINK \l "_ENREF_70" \o "Haussmann, 2012 #34578" 70]. Exposure to dexamethasone a potent synthetic GCs (That induced comparable effects to CORT) retarded growth rat through the inhibition of protein biosynthesis in Japanese quail embryos  ADDIN EN.CITE <EndNote><Cite><Author>Kaltner</Author><Year>1993</Year><RecNum>49977</RecNum><DisplayText>[79]</DisplayText><record><rec-number>49977</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">49977</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kaltner, H.</author><author>Schrott, M.</author><author>Schmahl, W.</author><author>Wittmann, J.</author></authors></contributors><auth-address>Institute of Physiology, Physiological Chemistry and Nutrition Physiology, Veterinary Faculty, Ludwig-Maximilians-University of Munich, Federal Republic of Germany.</auth-address><titles><title>Developmental retardation of the Japanese quail embryo under the influence of dexamethasone</title><secondary-title>Res Commun Chem Pathol Pharmacol</secondary-title></titles><periodical><full-title>Res Commun Chem Pathol Pharmacol</full-title></periodical><pages>259-73</pages><volume>79</volume><number>3</number><edition>1993/03/01</edition><keywords><keyword>Animals</keyword><keyword>Coturnix/*embryology</keyword><keyword>Dexamethasone/*toxicity</keyword><keyword>Embryo, Nonmammalian/drug effects</keyword><keyword>Embryonic Development</keyword><keyword>Kidney/drug effects/embryology</keyword></keywords><dates><year>1993</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0034-5164 (Print)&#xD;0034-5164 (Linking)</isbn><accession-num>8480074</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8480074</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_79" \o "Kaltner, 1993 #49977" 79] in addition inhibition of growth hormone releases from pituitary cultured cells  ADDIN EN.CITE <EndNote><Cite><Author>Tonshoff</Author><Year>1997</Year><RecNum>49974</RecNum><DisplayText>[80]</DisplayText><record><rec-number>49974</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">49974</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Tonshoff, B., and O. Mehls, </author></authors></contributors><titles><title> Interactions between glucocorticoids and the growth hormone-insulin-like growth factor&#xD;axis.</title><secondary-title> Pediatr. Transplant. </secondary-title></titles><volume>1:183-189.</volume><dates><year>1997</year></dates><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/10084779</url></related-urls></urls></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_80" \o "Tonshoff, 1997 #49974" 80]. In ovo CORT treatment elevated the circulating levels of CORT in hatch time in chickens  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_81" \o "Rodricks, 2006 #756" 81] and at 8 weeks old in Japanese quail  ADDIN EN.CITE <EndNote><Cite><Author>Hayward</Author><Year>2006</Year><RecNum>2466</RecNum><DisplayText>[82]</DisplayText><record><rec-number>2466</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">2466</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hayward, L. S.</author><author>Richardson, J. B.</author><author>Grogan, M. N.</author><author>Wingfield, J. C.</author></authors></contributors><auth-address>Department of Biology, University of Washington, Seattle, WA 98195, USA. lhayward@u.washington.edu</auth-address><titles><title>Sex differences in the organizational effects of corticosterone in the egg yolk of quail</title><secondary-title>Gen Comp Endocrinol</secondary-title></titles><periodical><full-title>Gen Comp Endocrinol</full-title></periodical><pages>144-8</pages><volume>146</volume><number>2</number><edition>2006/01/25</edition><keywords><keyword>Animals</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>Coturnix/embryology/*growth &amp; development</keyword><keyword>Egg Yolk/*chemistry</keyword><keyword>Female</keyword><keyword>Male</keyword><keyword>Sex Factors</keyword></keywords><dates><year>2006</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>0016-6480 (Print)&#xD;0016-6480 (Linking)</isbn><accession-num>16430891</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=16430891</url></related-urls></urls><electronic-resource-num>S0016-6480(05)00332-1 [pii]&#xD;10.1016/j.ygcen.2005.10.016</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_82" \o "Hayward, 2006 #2466" 82]. Similar results were reported in rats, revealing that exposure of CORT during gestation period inhibited fetal growth rate in mice  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_83" \o "Audette, 2011 #49501" 83,  HYPERLINK \l "_ENREF_84" \o "Vaughan, 2012 #49502" 84] and permanently increased the plasma CORT levels in adult rats  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_85" \o "Levitt, 1996 #42758" 85,  HYPERLINK \l "_ENREF_86" \o "Welberg, 2001 #46070" 86].
    Prenatal stress has been found to promote aggressive behavior in humans  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_87" \o "Tardiff, 1992 #46487" 87,  HYPERLINK \l "_ENREF_88" \o "Glover, 2011 #49827" 88]. In European starlings, the artificial elevation of yolk CORT enhanced offspring flight behaviors performance  ADDIN EN.CITE <EndNote><Cite><Author>Chin</Author><Year>2009</Year><RecNum>37318</RecNum><DisplayText>[18]</DisplayText><record><rec-number>37318</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37318</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Chin, E. H.</author><author>Love, O. P.</author><author>Verspoor, J. J.</author><author>Williams, T. D.</author><author>Rowley, K.</author><author>Burness, G.</author></authors></contributors><auth-address>Environmental and Life Sciences Graduate Program, Trent University, Peterborough, Ontario, Canada K9J 7B8.</auth-address><titles><title>Juveniles exposed to embryonic corticosterone have enhanced flight performance</title><secondary-title>Proc Biol Sci</secondary-title></titles><periodical><full-title>Proc Biol Sci</full-title></periodical><pages>499-505</pages><volume>276</volume><number>1656</number><edition>2008/10/10</edition><keywords><keyword>Animals</keyword><keyword>*Corticosterone</keyword><keyword>Embryo, Nonmammalian</keyword><keyword>*Flight, Animal</keyword><keyword>Starlings/*physiology</keyword><keyword>Stress, Physiological</keyword></keywords><dates><year>2009</year><pub-dates><date>Feb 7</date></pub-dates></dates><isbn>0962-8452 (Print)&#xD;0962-8452 (Linking)</isbn><accession-num>18842541</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=18842541</url></related-urls></urls><custom2>2664354</custom2><electronic-resource-num>F6H80G1662207650 [pii]&#xD;10.1098/rspb.2008.1294</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_18" \o "Chin, 2009 #37318" 18], increased fearfulness behavior in chickens  ADDIN EN.CITE <EndNote><Cite><Author>Janczak</Author><Year>2006</Year><RecNum>37863</RecNum><DisplayText>[14]</DisplayText><record><rec-number>37863</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37863</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Janczak, A. M.</author><author>Braastad, B. O.</author><author>Bakken, M.</author></authors></contributors><auth-address>Janczak, AM&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, Box 5003, N-1432 As, Norway&#xD;Norwegian Univ Life Sci, Dept Anim &amp; Aquacultural Sci, N-1432 As, Norway</auth-address><titles><title>Behavioural effects of embryonic exposure to corticosterone in chickens</title><secondary-title>Applied Animal Behaviour Science</secondary-title><alt-title>Appl Anim Behav Sci</alt-title></titles><periodical><full-title>Applied Animal Behaviour Science</full-title></periodical><alt-periodical><full-title>Appl Anim Behav Sci</full-title></alt-periodical><pages>69-82</pages><volume>96</volume><number>1-2</number><keywords><keyword>prenatal stress</keyword><keyword>chicken</keyword><keyword>fear</keyword><keyword>cognition</keyword><keyword>competition</keyword><keyword>prenatal stress</keyword><keyword>feather pecking</keyword><keyword>plasma-corticosterone</keyword><keyword>pregnancy</keyword><keyword>animals</keyword><keyword>anxiety</keyword><keyword>alters</keyword><keyword>yolk</keyword><keyword>rats</keyword><keyword>hen</keyword></keywords><dates><year>2006</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0168-1591</isbn><accession-num>ISI:000235262600006</accession-num><urls><related-urls><url>&lt;Go to ISI&gt;://000235262600006</url></related-urls></urls><electronic-resource-num>DOI 10.1016/j.applanim.2005.04.020</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_14" \o "Janczak, 2006 #37863" 14], enhanced recall of a passive avoidance task in chickens  ADDIN EN.CITE <EndNote><Cite><Author>Sui</Author><Year>1997</Year><RecNum>1722</RecNum><DisplayText>[19]</DisplayText><record><rec-number>1722</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">1722</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sui, N.</author><author>Sandi, C.</author><author>Rose, S. P.</author></authors></contributors><auth-address>Brain and Behaviour Centre, Academia Sinica, Beijing, People&apos;s Republic of China. suinan@ihw.co.cn</auth-address><titles><title>Interactions of corticosterone and embryonic light deprivation on memory retention in day-old chicks</title><secondary-title>Brain Res Dev Brain Res</secondary-title></titles><periodical><full-title>Brain Res Dev Brain Res</full-title></periodical><pages>269-72</pages><volume>101</volume><number>1-2</number><edition>1997/07/18</edition><keywords><keyword>Animals</keyword><keyword>Anti-Inflammatory Agents/*pharmacology</keyword><keyword>Avoidance Learning/drug effects/physiology</keyword><keyword>Chick Embryo</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>*Darkness</keyword><keyword>Memory/*drug effects/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Jul 18</date></pub-dates></dates><isbn>0165-3806 (Print)&#xD;0165-3806 (Linking)</isbn><accession-num>9263600</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9263600</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_19" \o "Sui, 1997 #1722" 19], and increased the pecking behavior and pebbles in chickens  ADDIN EN.CITE <EndNote><Cite><Author>Freire</Author><Year>2006</Year><RecNum>37754</RecNum><DisplayText>[20]</DisplayText><record><rec-number>37754</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">37754</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Freire, R.</author><author>van Dort, S.</author><author>Rogers, L. J.</author></authors></contributors><auth-address>Centre for Neuroscience and Animal Behaviour, University of New England, Australia. rfreire2@une.edu.au</auth-address><titles><title>Pre- and post-hatching effects of corticosterone treatment on behavior of the domestic chick</title><secondary-title>Horm Behav</secondary-title></titles><periodical><full-title>Horm Behav</full-title></periodical><pages>157-65</pages><volume>49</volume><number>2</number><edition>2005/07/02</edition><keywords><keyword>Animals</keyword><keyword>Animals, Newborn/*physiology</keyword><keyword>Behavior, Animal/*drug effects</keyword><keyword>Chick Embryo</keyword><keyword>Chickens/*physiology</keyword><keyword>Corticosterone/*pharmacology</keyword><keyword>Female</keyword><keyword>Functional Laterality/physiology</keyword><keyword>Hawks</keyword><keyword>Male</keyword><keyword>Photic Stimulation</keyword><keyword>Predatory Behavior</keyword><keyword>Psychomotor Performance/drug effects</keyword></keywords><dates><year>2006</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0018-506X (Print)&#xD;0018-506X (Linking)</isbn><accession-num>15990098</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15990098</url></related-urls></urls><electronic-resource-num>S0018-506X(05)00143-1 [pii]&#xD;10.1016/j.yhbeh.2005.05.015</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_20" \o "Freire, 2006 #37754" 20]. In seabirds, the implanted CORT caused increases in the aggressive behavior when compared to the controls  ADDIN EN.CITE <EndNote><Cite><Author>Kitaysky</Author><Year>2003</Year><RecNum>33612</RecNum><DisplayText>[89]</DisplayText><record><rec-number>33612</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">33612</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kitaysky, A. S.</author><author>Kitaiskaia, E. V.</author><author>Piatt, J. F.</author><author>Wingfield, J. C.</author></authors></contributors><auth-address>Department of Zoology, 24 Kincaid Hall, Box 351800, University of Washington, Seattle, WA 98195, USA. kitaysky@u.washington.edu</auth-address><titles><title>Benefits and costs of increased levels of corticosterone in seabird chicks</title><secondary-title>Horm Behav</secondary-title></titles><periodical><full-title>Horm Behav</full-title></periodical><pages>140-9</pages><volume>43</volume><number>1</number><edition>2003/03/05</edition><keywords><keyword>Aggression/drug effects/*physiology</keyword><keyword>Animals</keyword><keyword>Association Learning/drug effects/*physiology</keyword><keyword>Birds/growth &amp; development/*physiology</keyword><keyword>Cognition/drug effects/physiology</keyword><keyword>Corticosterone/*blood/pharmacology</keyword><keyword>Drug Implants</keyword><keyword>Eating/drug effects/physiology</keyword><keyword>Environment</keyword><keyword>Feeding Behavior/drug effects/*physiology</keyword><keyword>Memory/drug effects/physiology</keyword><keyword>Space Perception/drug effects/physiology</keyword><keyword>Stress, Physiological/*physiopathology</keyword></keywords><dates><year>2003</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0018-506X (Print)&#xD;0018-506X (Linking)</isbn><accession-num>12614644</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12614644</url></related-urls></urls><electronic-resource-num>S0018506X02000302 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_89" \o "Kitaysky, 2003 #33612" 89]. In agreement with these results, we found increased frequency of tonic immobility and aggressive behaviors in chickens that prenatally exposure to the high dose of CORT. 
    Prenatal stress during late gestational period causes alterations in 11�-HSD1 mRNA expression in marmosets�  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_90" \o "Nyirenda, 2009 #7396" 90], human  ADDIN EN.CITE <EndNote><Cite><Author>Baud</Author><Year>2019</Year><RecNum>67274</RecNum><DisplayText>[91]</DisplayText><record><rec-number>67274</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67274</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Baud,Olivier</author><author>Berkane,Nadia</author></authors></contributors><titles><title>Hormonal Changes Associated With Intra-Uterine Growth Restriction: Impact on the Developing Brain and Future Neurodevelopment</title><secondary-title>Frontiers in Endocrinology</secondary-title><short-title>Intra-uterine growth restriction and hormones</short-title></titles><periodical><full-title>Frontiers in endocrinology</full-title></periodical><volume>10</volume><number>179</number><keywords><keyword>Intra-uterine growth restriction,Glucocorticosteroids,Neurosteroids,Thyroid Hormones,Insulin-Growth Factor Type-1</keyword></keywords><dates><year>2019</year><pub-dates><date>2019-March-26</date></pub-dates></dates><isbn>1664-2392</isbn><work-type>Review</work-type><urls><related-urls><url>https://www.frontiersin.org/article/10.3389/fendo.2019.00179</url></related-urls></urls><electronic-resource-num>10.3389/fendo.2019.00179</electronic-resource-num><language>English</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_91" \o "Baud, 2019 #67274" 91], rodents  ADDIN EN.CITE <EndNote><Cite><Author>Christoforou</Author><Year>2020</Year><RecNum>67275</RecNum><DisplayText>[92]</DisplayText><record><rec-number>67275</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67275</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Christoforou, Efthimia R.</author><author>Sferruzzi-Perri, Amanda N.</author></authors></contributors><titles><title>Molecular mechanisms governing offspring metabolic programming in rodent models of in utero stress</title><secondary-title>Cellular and Molecular Life Sciences</secondary-title></titles><periodical><full-title>Cellular and Molecular Life Sciences</full-title></periodical><dates><year>2020</year><pub-dates><date>2020/06/03</date></pub-dates></dates><isbn>1420-9071</isbn><urls><related-urls><url>https://doi.org/10.1007/s00018-020-03566-z</url></related-urls></urls><electronic-resource-num>10.1007/s00018-020-03566-z</electronic-resource-num></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_92" \o "Christoforou, 2020 #67275" 92], and decreased 11�-HSD2 mRNA expression in later life in mammals (Review)  ADDIN EN.CITE <EndNote><Cite><Author>Charil</Author><Year>2010</Year><RecNum>49964</RecNum><DisplayText>[93]</DisplayText><record><rec-number>49964</rec-number><foreign-keys><key app="EN" db-id="ssxdpzezqvdww8exxvxprdsu2xxzftdxfwww">49964</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Charil, A.</author><author>Laplante, D. P.</author><author>Vaillancourt, C.</author><author>King, S.</author></authors></contributors><auth-address>McGill University, Department of Psychiatry, Montreal, Quebec, Canada.</auth-address><titles><title>Prenatal stress and brain development</title><secondary-title>Brain Res Rev</secondary-title></titles><pages>56-79</pages><volume>65</volume><number>1</number><edition>2010/06/17</edition><keywords><keyword>Adult</keyword><keyword>Animals</keyword><keyword>Brain/*growth &amp; development/physiology</keyword><keyword>Disasters</keyword><keyword>Female</keyword><keyword>Fetus/physiology</keyword><keyword>Humans</keyword><keyword>Hydrocortisone/physiology</keyword><keyword>Hypothalamo-Hypophyseal System/embryology</keyword><keyword>Magnetic Resonance Imaging</keyword><keyword>Male</keyword><keyword>Pituitary-Adrenal System/embryology</keyword><keyword>Pregnancy</keyword><keyword>Pregnancy, Animal/physiology</keyword><keyword>Prenatal Exposure Delayed Effects/*pathology</keyword><keyword>Sex Characteristics</keyword><keyword>Stress, Psychological/complications/*pathology</keyword></keywords><dates><year>2010</year><pub-dates><date>Oct 5</date></pub-dates></dates><isbn>1872-6321 (Electronic)&#xD;0165-0173 (Linking)</isbn><accession-num>20550950</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=20550950</url></related-urls></urls><electronic-resource-num>S0165-0173(10)00072-X [pii]&#xD;10.1016/j.brainresrev.2010.06.002</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_93" \o "Charil, 2010 #49964" 93]. In agreement with those findings, High dose administration of CORT upregulated hypothalamic 11�-HSD1 expression whereas, downregulated 11�-HSD2 mRNA in the hypothalamus. In mammalian species, upregulation of hippocampal 11�-HSD1 mRNA expression together with cerebral cortex resulted in decline of cognitive which was associated with aging in mice  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_94" \o "Holmes, 1997 #50557" 94]. In humans and rodents, 11�-HSD1 over expression was associated with obesity in later life  ADDIN EN.CITE <EndNote><Cite><Author>Livingstone</Author><Year>2000</Year><RecNum>50558</RecNum><DisplayText>[95]</DisplayText><record><rec-number>50558</rec-number><foreign-keys><key app="EN" db-id="ssxdpzezqvdww8exxvxprdsu2xxzftdxfwww">50558</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Livingstone, D. E.</author><author>Jones, G. C.</author><author>Smith, K.</author><author>Jamieson, P. M.</author><author>Andrew, R.</author><author>Kenyon, C. J.</author><author>Walker, B. R.</author></authors></contributors><auth-address>Department of Medical Sciences, University of Edinburgh, Western General Hospital, United Kingdom.</auth-address><titles><title>Understanding the role of glucocorticoids in obesity: tissue-specific alterations of corticosterone metabolism in obese Zucker rats</title><secondary-title>Endocrinology</secondary-title></titles><pages>560-3</pages><volume>141</volume><number>2</number><edition>2000/01/29</edition><keywords><keyword>11-beta-Hydroxysteroid Dehydrogenase Type 1</keyword><keyword>3-Oxo-5-alpha-Steroid 4-Dehydrogenase/metabolism</keyword><keyword>Adrenal Cortex Hormones/blood/*metabolism/urine</keyword><keyword>Animals</keyword><keyword>Corticosterone/analogs &amp; derivatives/*metabolism</keyword><keyword>Humans</keyword><keyword>Hydroxysteroid Dehydrogenases/genetics/metabolism</keyword><keyword>Liver/enzymology</keyword><keyword>Male</keyword><keyword>Microsomes, Liver/enzymology</keyword><keyword>Obesity/genetics/*metabolism</keyword><keyword>Organ Specificity</keyword><keyword>Rats</keyword><keyword>Rats, Zucker</keyword><keyword>Thinness</keyword></keywords><dates><year>2000</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0013-7227 (Print)&#xD;0013-7227 (Linking)</isbn><accession-num>10650936</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10650936</url></related-urls></urls><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_95" \o "Livingstone, 2000 #50558" 95]. The reduction in the activity or expression of 11�-HSD2 during pregnancy was resulted in development of metabolic syndromes  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_96" \o "Ni, 2018 #67276" 96]   such as hypertension  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_97" \o "Kosicka, 2016 #67277" 97,  HYPERLINK \l "_ENREF_98" \o "Kosicka, 2018 #67279" 98], glucose intolerance  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_99" \o "Wyrwoll, 2009 #67278" 99]as well as the programming of HPA axis activity  ADDIN EN.CITE <EndNote><Cite><Author>McGowan</Author><Year>2017</Year><RecNum>67280</RecNum><DisplayText>[100]</DisplayText><record><rec-number>67280</rec-number><foreign-keys><key app="EN" db-id="vrxdrtefjtswfoesr27vev90a2att20ztrdd">67280</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>McGowan, Patrick O</author><author>Matthews, Stephen G</author></authors></contributors><titles><title>Prenatal Stress, Glucocorticoids, and Developmental Programming of the Stress Response</title><secondary-title>Endocrinology</secondary-title></titles><periodical><full-title>Endocrinology</full-title><abbr-1>Endocrinology</abbr-1></periodical><pages>69-82</pages><volume>159</volume><number>1</number><dates><year>2017</year></dates><isbn>0013-7227</isbn><urls><related-urls><url>https://doi.org/10.1210/en.2017-00896</url></related-urls></urls><electronic-resource-num>10.1210/en.2017-00896</electronic-resource-num><access-date>9/2/2020</access-date></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_100" \o "McGowan, 2017 #67280" 100]and anxiety related behaviors in later life  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_42" \o "Welberg, 2000 #15594" 42,  HYPERLINK \l "_ENREF_101" \o "Edwards, 1993 #24810" 101,  HYPERLINK \l "_ENREF_102" \o "Harris, 2011 #48480" 102]. 
In this study, the increases of aggressive behaviors in exposed high CORT chickens were associated with alterations in the hypothalamic expression of serotonergic genes mRNA. A significant increase was found in hypothalamic 5HTR1A mRNA expression in response to CORT treatment. There are several lines of evidence indicating that the neurocircuits for stress and aggression are reciprocally interrelated in non mammalian species  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_103" \o "Summers, 2005 #50611" 103-105]. Animals predisposed to be dominant often show higher GCs associated with lower 5-HT  ADDIN EN.CITE <EndNote><Cite><Author>Summers</Author><Year>2006</Year><RecNum>50610</RecNum><DisplayText>[46]</DisplayText><record><rec-number>50610</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">50610</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Summers, C. H.</author><author>Winberg, S.</author></authors></contributors><auth-address>Department of Biology, University of South Dakota, Vermillion, SD 57069 USA. cliff@usd.edu</auth-address><titles><title>Interactions between the neural regulation of stress and aggression</title><secondary-title>J Exp Biol</secondary-title></titles><periodical><full-title>J Exp Biol</full-title></periodical><pages>4581-9</pages><volume>209</volume><number>Pt 23</number><edition>2006/11/23</edition><keywords><keyword>Aggression/*physiology</keyword><keyword>Animals</keyword><keyword>Neurons/*physiology</keyword><keyword>Stress, Physiological/*physiopathology</keyword></keywords><dates><year>2006</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>0022-0949 (Print)&#xD;0022-0949 (Linking)</isbn><accession-num>17114393</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=17114393</url></related-urls></urls><electronic-resource-num>209/23/4581 [pii]&#xD;10.1242/jeb.02565</electronic-resource-num><language>eng</language></record></Cite></EndNote>[ HYPERLINK \l "_ENREF_46" \o "Summers, 2006 #50610" 46]. GCs and prenatal stress were reported to enhance 5-HT system function and increase 5-HTR1A mRNA expression in rat  ADDIN EN.CITE  ADDIN EN.CITE.DATA [ HYPERLINK \l "_ENREF_106" \o "Wang, 2012 #50648" 106,  HYPERLINK \l "_ENREF_107" \o "Goodfellow, 2009 #15541" 107], which seems to agree with our current finding. However, it is remain unknown whether these effects will�be�transmitted�to�the next generation in chickens, thus, the stressed mother that deposited high amounts CORT in the eggs may have similar phenotypic and behavioral outcomes.� 

Conclusion
    In conclusion, our findings suggest that prenatal CORT exposure may influence the phenotype, aggressive behavior and tonic immobility of chickens. Changes in hypothalamic GCs metabolic enzymes and 5-HT system gene expression could be due to the consequence of earlier effects on growth retardation. Further studies are required to clarify the transgenerational effect of CORT in ovo in chickens.
 
Acknowledgments
This work was supported by the NSFC-Guangdong Joint Fund (Project No. U0931004), the Special Fund for Agro-scientific Research in the Public Interest (201003011), and the Priority Academic Program Development of Jiangsu Higher Education Institutions. Especial thanks conducted to Professor Donald C. Lay his critical revision of manuscript.

Conflict of interest
No conflict of interests exists.
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Figure legends
    Figure 1. (A) Effect of in ovo injection of CORT on chicken post hatch growth performance. (B) Effect of in ovo injection of CORT on chicken aggressive behavior (grabbing, nape each other, pecking and twisting) at week 25 and (C) Effect of in ovo injection of CORT on TI test. (D) Effect of in ovo injection of CORT on plasma CORT concentrations. The values are the means � SEM, n = 25-30/group for the growth curve, n= 25-30/group for the behavioral analysis, TI and = 6 for plasma CORT concentration. 
    Figure 2. (A) Effect of in ovo injection of CORT on hypothalamic 11�-HSD1 mRNA, (B) 11�-HSD2 mRNA, (C) 20-HSD mRNA, (D) GR mRNA and (E) MR mRNA expression. Values are the mean � SEM, n = 6/group. Bars with different letters are significantly different at P<0.05, for 11�-HSD1, 11�-HSD2 and MR 
Figure 3. (A) Effect of in ovo injection of CORT on hypothalamic 5-HTR1A mRNA and (B) HTR1B mRNA expression. Values are the mean � SEM, n = 6/group. Bars with different letters are significantly different at P<0.05, for 5-HTR1A 

Fig. 1
A



B


C


D

E


Fig. 2
A



B


C

D


E



Fig. 3

A


B


Table 1 Real-Time PCR Primers
Target genesGen Bank accession numberPCR products (bp)Primer sequences#Z[�[�\;]^}^�^N_`�`�a�b�cd�dce�frg2h�h�i�j8m�np�pZq���������������������������d�gd�YI	d��gd�YIZq!r�r�s't�t�u�v�v�v�vTy�{�|�|�|�|�|�|�|�������������������d�gd��d�gd��d�gd�]*d�7$8$H$gd�]*$d�7$8$H$a$gdS{�d�gd�P�
��gd�E�d�gd�YI	d��gd�YI�v�v�v�v�vww"w�w�w�w�w�w�w�w�w�w�wx�x�x�x�����������fMf1h(9hS{�CJOJQJ^JaJfHq�
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����"h(9h�[�6�CJOJQJ]�^Jh(9h�[�CJOJQJ^Jh(9h�X�CJOJQJ^J"h(9h�P�6�CJOJQJ]�^J"h(9h_%g6�CJOJQJ]�^Jh(9h_%gCJOJQJ^Jh(9h�P�CJOJQJ^Jh(9h^6CJOJQJ^J�x�xyyyyNyPyRyTy\yjylypyxy�y�y��ī����yj[L[=[+"h(9h�P�6�CJOJQJ]�^Jh(9h�[�CJOJQJ^Jh(9h�X�CJOJQJ^Jh(9h�P�CJOJQJ^Jh(9h^6CJOJQJ^J1h(9h�P�CJOJQJ^JaJfHq�
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CJOJQJ^J!j(�h(9h�*8OJQJU^Jh(9h�OJQJ^J!je�h(9hYm OJQJU^Jh(9h�;�OJQJ^J!jl�h(9h#i�OJQJU^Jh(9h��OJQJ^Jh(9h-
?OJQJ^Jh(9h�lIOJQJ^J!jr�h(9h%>OJQJU^J,j��h(9h�c�OJQJU^JmHnHuh(9h�c�OJQJ^Jh(9h<
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}}}+}~�l����µ����}�p�_�N=;=U h
$h��CJOJQJ^JaJ h(9h��CJOJQJ^JaJ!j��h(9h9AOJQJU^Jh(9h�,lOJQJ^J!j��h(9h9AOJQJU^Jh(9h9AOJQJ^Jh(9hD�OJQJ^Jh(9h��OJQJ^Jh(9h/]OJQJ^Jh(9hYm CJOJQJ^J!j��h(9hYm OJQJU^Jh(9h�*8CJOJQJ^Jh(9h-
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la��-actinL08165300F : 52 - TGCGTGACATCAAGGAGAAG -32 
R : 52 - TGCCAGGGTACATTGTGGTA -32 GR DQ227738102F : 52 - CTTCCATCCGCCCTTCA -32 
R : 52 - TCGCATCTGTTTCACCC -32 MRNM_001159345.1210F: 52 - ACGCAGGATATGACAGCTCG-32 
R: 52 - AGTACAGGGGCTTGGCATTC-32 11�-HSD1 HYPERLINK "http://www.ncbi.nlm.nih.gov/nucleotide/118102456?report=genbank&log$=nucltop&blast_rank=1&RID=73C6S6TB014" \o "Show report for XM_417988.2" XM_417988.2
229F: 52 -GGTGGTGAAAGAGGCTGAGAAC-32 
R: 52 -GGAGGCGACTTTACCTGAAACAG-32 11�-HSD2 HYPERLINK "http://www.ncbi.nlm.nih.gov/nucleotide/326927090?report=genbank&log$=nucltop&blast_rank=1&RID=73CGHNZR01S" \o "Show report for XM_003209680.1" XM_003209680.1229F: 52 -GGTGGTGAAAGAGGCTGAGAACA-32 
R: 52 -GGAGGCGACTTTACCTGAAACAG-32 20-HSD HYPERLINK "http://www.ncbi.nlm.nih.gov/nucleotide/71895266?report=genbank&log$=nucltop&blast_rank=1&RID=73CKZM1Y015" \o "Show report for NM_001030795.1" NM_001030795.1220F: 52 - CATCCTGAGAAGATAATGTCCAACG -32 
R: 52 - TGCTTTGCAGATCATCAATATCCAG -32 5-HT1AGU189388.1202F: 52 - AGAACACGGAGGCCAAGC -32 
R: 52 - ACGGCAACCAGCAGAGGA -32 5-HT1BGU385013.1133F: 52 - CACGGACCACGTCCTCTACAC -32 
R: 52 - TTTCTTTGGCGTCTGCTTCA -32 








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