Chicken as a Developmental Model


The development of a complex organism from a single cell, the fertilised egg, has fascinated people for centuries. Embryo development is highly reproducible and exquisitely regulated. How is it that all tissues and organs form in the right places and at the right time? How is the development of different organ systems coordinated, so that they all fit together correctly at the end? It is challenging to study development, because many embryos are small or inaccessible. The chick embryo is a popular model system with many experimental advantages, which include classic ‘cut and paste’ experiments and mechanistic gene function analyses. The combination of micromanipulations with gain‐ or loss‐of‐function is particularly powerful. The recent development of transgenic lines and advanced imaging techniques ensure that the chicken remains an attractive model system, which will continue to make major contributions to our understanding of molecular and cellular mechanisms controlling developmental processes.

Key Concepts

  • The chick embryo is easy to access and observe and is therefore an attractive model to study developmental processes.
  • In the early stages, chick embryo morphology is very similar to human, both are amniotes and their development is very similar.
  • The chicken and human genomes share considerable homology.
  • Classic embryological manipulations, such as tissue ablations and tissue grafts, provide information about the contribution of particular cell populations to various organs. These experiments have also told us how cells and tissues influence each other in their developmental decisions.
  • The molecular and cellular basis for many developmental processes and phenomena were first described in the chick, including limb patterning, neural crest migration, dorso‐ventral neural tube patterning, blood vessel formation, somite segmentation and left–right asymmetry.
  • Experiments in chicken, which examine the function of genes, have helped elucidate the underlying mechanisms of human genetic diseases and provide a basis for testing novel therapies.
  • Time‐lapse video microscopy can be used to image live chick embryos, either in ovo or ex ovo using embryo culture.
  • We can use chick embryos to visualise complex processes, including cell migration, cell–cell communication, cell differentiation and tissue morphogenesis, in an amniote system.

Keywords: chick; quail; vertebrate; tissue graft; microsurgery; dye injection; electroporation; embryo culture; live imaging

Figure 1. Bead implantation and detection of gene expression patterns by hybridisation or immunostaining. (a) Beads soaked in growth factors or pharmacological inhibitors or activators of signalling pathways can be implanted into developing embryos. A bead implanted into the forelimb is shown. (b) Detection of specific messenger RNA transcripts in whole‐mount embryos with anti‐sense RNA probes against Fgf‐8 (b), Myogenin = Mgn (c), or ventricular myosin heavy chain = vMHC (d). Probes incorporate a DIG‐UTP nucleotide, which is detected using an alkaline phosphatase‐coupled anti‐DIG antibody. Alkaline phosphatase enzyme converts a substrate into a coloured precipitate, which generates a localised signal that is easily detected. Different structures are indicated by arrows in the different panels, AER = apical ectodermal ridge of the limb bud, so = somite, ht = heart. Other structures also expressing these genes are not indicated. (e) Protein can be detected by antibody staining in whole mount; the fluorescent signal indicates localisation of vMHC.
Figure 2. Microinjection and electroporation of chicken embryos in EC culture or . (a) The injection set‐up consists of a stereo‐dissection microscope, micromanipulators, a pressure injector, an electroporator and a light source. (b) Close‐up of the EC‐culture dish, which contains the embryo mounted on a filter paper frame and placed on a semi‐solid medium. (c) Close‐up of the HH3 embryo in the filter paper carrier, illustrating the placement of the electrodes on either side of the primitive streak, indicated by a stippled line. (d) An embryo, which was electroporated at HH3 with a GFP‐plasmid and incubated to HH5. The GFP‐expressing cells have migrated away from the site of injection (bright green) and are now distributed in an arc shape. Many of these cells are prospective cardiac cells and will migrate to form the heart. (e) Electroporation of an embryo . Black ink injected beneath the embryo helps visualise it. (f) A chick embryo specifically expressing GFP in one half of the neural tube (nt), other structures indicated are the eye (ey) and the heart (ht), which is filled with red blood cells. (g) Section through the embryo shown in (f) shows restricted expression of the electroporated GFP plasmid on one side of the neural tube (nt).
Figure 3. Fate mapping using GFP‐transgenic chicken embryos. (a) A single somite, micro‐dissected from a GFP‐transgenic embryo, was grafted into a non‐transgenic host embryo (white arrow in b). (c) After 7 days of incubation, the somite has contributed to the vertebral column in the neck region, as indicated by the presence of GFP‐positive cells. (d) More detailed analysis shows that a single somite contributes cells (purple colour) to two different vertebrae, which are indicated by a stippled outline. The phenomenon, whereby the original segments (=somites) contribute to the posterior and anterior half of new segments (=vertebrae) is called ‘re‐segmentation’. (Courtesy of Mike McGrew, Roslin Institute, Edinburgh.)
Figure 4. Transgenic chick embryos expressing fluorescent markers, mApple (red) or eGFP (green), specifically in macrophages, under the control of the CSF1R promoter. (a) Macrophages (red) are distributed throughout the developing embryo. (b,c) Macrophages in limbs (green) at early (b) or later (c) stages of development. In (c), the limb has been co‐stained with Lysotracker, a chemical that is picked up by active macrophages that are cleaning up apoptotic cells between the forming digits. These macrophages therefore appear yellow. (d) Distribution of macrophages (red) in skeletal muscle tissue. (Courtesy of Adam Balic, David Hume and Helen Sang, Roslin Institute, Edinburgh; see also Balic ., .)


Abu‐Elmagd M, Robson L, Sweetman D, et al. (2010) Wnt/Lef1 signaling acts via Pitx2 to regulate somite myogenesis. Developmental Biology 337 (2): 211–219.

Ainsworth SJ, Stanley RL, Evans DJ (2010) Developmental stages of the Japanese quail. Journal of Anatomy 216 (1): 3–15. DOI: 10.1111/j.1469-7580.2009.01173.x.

Balic A, Garcia‐Morales C, Vervelde L, et al. (2014) Visualisation of chicken macrophages using transgenic reporter genes: insights into the development of the avian macrophage lineage. Development 141 (16): 3255–3265.

Bangs F, Antonio N, Thongnuek P, et al. (2011) Generation of mice with functional inactivation of talpid3, a gene first identified in chicken. Development 138 (15): 3261–3272.

Becker DL, McGonnell I, Makarenkova HP, et al. (1999) Roles for alpha 1 connexin in morphogenesis of chick embryos revealed using a novel antisense approach. Developmental Genetics 24 (1–2): 33–42.

Bellairs R, Lorenz FW and Dunlap T (1978) Cleavage in the chick embryo. Journal of Embryology and Experimental Morphology 43: 55–69.

Chapman SC, Collignon J, Schoenwolf GC and Lumsden A (2001) Improved method for chick whole‐embryo culture using a filter paper carrier. Developmental Dynamics 220 (3): 284–289.

Chapman SC, Lawson A, Macarthur WC, et al. (2005) Ubiquitous GFP expression in transgenic chickens using a lentiviral vector. Development 132 (5): 935–940.

Christ B, Huang R and Scaal M (2004) Formation and differentiation of the avian sclerotome. Anatomy and Embryology (Berlin) 208 (5): 333–350.

Chuai M, Dormann D and Weijer CJ (2009) Imaging cell signalling and movement in development. Seminars in Cell & Developmental Biology 20 (8): 947–955.

Chuai M, Hughes D and Weijer CJ (2012) Collective epithelial and mesenchymal cell migration during gastrulation. Current Genomics 13 (4): 267–277.

Chuai M, Zeng W, Yang X, et al. (2006) Cell movement during chick primitive streak formation. Developmental Biology 296 (1): 137–149.

Clinton M, Zhao D, Nandi S and McBride D (2012) Evidence for avian cell autonomous sex identity (CASI) and implications for the sex‐determination process? Chromosome Research: An International Journal on the Molecular, Supramolecular and Evolutionary Aspects of Chromosome Biology 20 (1): 177–190.

Consortium, I. C. G. S (2004) Sequence and comparative analysis of the chicken genome provide unique perspectives on vertebrate evolution. Nature 432 (7018): 695–716.

Davey MG, Paton IR, Yin Y, et al. (2006) The chicken talpid3 gene encodes a novel protein essential for Hedgehog signaling. Genes & Development 20 (10): 1365–1377.

Eblaghie MC, Lunn JS, Dickinson RJ, et al. (2003) Negative feedback regulation of FGF signaling levels by Pyst1/MKP3 in chick embryos. Current Biology 13 (12): 1009–1018.

Eyal‐Giladi H and Kochav S (1976) From cleavage to primitive streak formation: a complementary normal table and a new look at the first stages of the development of the chick. Developmental Biology 49: 321–337.

Flentke GR, Garic A, Hernandez M and Smith SM (2014) CaMKII represses transcriptionally active beta‐catenin to mediate acute ethanol neurodegeneration and can phosphorylate beta‐catenin. Journal of Neurochemistry 128 (4): 523–535.

Goljanek‐Whysall K, Mok GF, Fahad Alrefaei A, et al. (2014) myomiR‐dependent switching of BAF60 variant incorporation into Brg1 chromatin remodeling complexes during embryo myogenesis. Development 141 (17): 3378–3387.

Griffin DK, Haberman F, Masabanda J, et al. (1999) Micro‐ and macrochromosome paints generated by flow cytometry and microdissection: tools for mapping the chicken genome. Cytogenetics and Cell Genetics 87 (3–4): 278–281.

Grocott T, Johnson S, Bailey AP and Streit A (2011) Neural crest cells organize the eye via TGF‐beta and canonical Wnt signalling. Nature Communications 2: 265.

Hamburger V and Hamilton HL (1951) A series of normal stages in the development of the chick embryo. Journal of Morphology 88 (1): 49–92.

Harpavat S and Cepko CL (2006) RCAS-RNAi: a loss-of-function method for the developing chick retina. BMC Developmental Biology 6: 2. DOI: 10.1186/1471-213X-6-2.

Hubbard SJ, Grafham DV, Beattie KJ, et al. (2005) Transcriptome analysis for the chicken based on 19,626 finished cDNA sequences and 485,337 expressed sequence tags. Genome Research 15 (1): 174–183.

Iimura T, Yang X, Weijer CJ and Pourquie O (2007) Dual mode of paraxial mesoderm formation during chick gastrulation. Proceedings of the National Academy of Sciences of the United States of America 104 (8): 2744–2749.

Itasaki N, Bel‐Vialar S and Krumlauf R (1999) Shocking' developments in chick embryology: electroporation and in ovo gene expression. Nature Cell Biology 1 (8): E203–E207.

Korn MJ and Cramer KS (2007) Windowing chicken eggs for developmental studies. Journal of Visualized Experiments 8: 306.

Le Douarin N, Dieterlen‐Lievre F, Creuzet S and Teillet MA (2008) Quail‐chick transplantations. Methods in Cell Biology 87: 19–58.

Lopez‐Sanchez C, Puelles L, Garcia‐Martinez V and Rodriguez‐Gallardo L (2005) Morphological and molecular analysis of the early developing chick requires an expanded series of primitive streak stages. Journal of Morphology 264 (1): 105–116.

Ma ZL, Wang G, Cheng X, et al. (2014) Excess caffeine exposure impairs eye development during chick embryogenesis. Journal of Cellular and Molecular Medicine 18 (6): 1134–1143.

Maroto M, Reshef R, Münsterberg AE, et al. (1997) Ectopic Pax‐3 activates MyoD and Myf‐5 expression in embryonic mesoderm and neural tissue. Cell 89 (1): 139–148.

Masabanda JS, Burt DW, O'Brien PC, et al. (2004) Molecular cytogenetic definition of the chicken genome: the first complete avian karyotype. Genetics 166 (3): 1367–1373.

McGrew MJ, Sherman A, Ellard FM, et al. (2004) Efficient production of germline transgenic chickens using lentiviral vectors. EMBO Reports 5 (7): 728–733.

McGrew MJ, Sherman A, Lillico SG, et al. (2008) Localised axial progenitor cell populations in the avian tail bud are not committed to a posterior Hox identity. Development 135 (13): 2289–2299.

Morgan BA and Fekete DM (1996) Manipulating gene expression with replication‐competent retroviruses. In: Bronner‐Fraser M, (ed.) Methods in Avian Embryology, vol. 51. San Diego, CA: Academic Press, Inc.

Nagai H, Sezaki M, Nakamura H and Sheng G (2014) Extending the limits of avian embryo culture with the modified Cornish pasty and whole‐embryo transplantation methods. Methods 66 (3): 441–446.

Norris A and Streit A (2014) Morpholinos: studying gene function in the chick. Methods 66 (3): 454–465.

Pekarik V, Bourikas D, Miglino N, et al. (2003) Screening for gene function in chicken embryo using RNAi and electroporation. Nature Biotechnology 21 (1): 93–96.

Pourquie O (2004) The chick embryo: a leading model in somitogenesis studies. Mechanisms of Development 121 (9): 1069–1079.

Poynter G, Huss D and Lansford R (2009) Japanese quail: an efficient animal model for the production of transgenic avians. Cold Spring Harbor Protocols 2009 (1): pdb emo112.

Rios AC, Denans N and Marcelle C (2010) Real‐time observation of Wnt beta‐catenin signaling in the chick embryo. Developmental Dynamics 239 (1): 346–353.

Rutland C, Warner L, Thorpe A, et al. (2009) Knockdown of alpha myosin heavy chain disrupts the cytoskeleton and leads to multiple defects during chick cardiogenesis. Journal of Anatomy 214 (6): 905–915.

Sanz‐Ezquerro JJ and Tickle C (2003) Fgf signaling controls the number of phalanges and tip formation in developing digits. Current Biology 13 (20): 1830–1836.

Sato Y, Kasai T, Nakagawa S, et al. (2007) Stable integration and conditional expression of electroporated transgenes in chicken embryos. Developmental Biology 305 (2): 616–624.

Sato Y, Poynter G, Huss D, et al. (2010) Dynamic analysis of vascular morphogenesis using transgenic quail embryos. PLoS One 5 (9): e12674.

Scaal M and Christ B (2004) Formation and differentiation of the avian dermomyotome. Anatomy and Embryology (Berlin) 208 (6): 411–424.

Scaal M, Gros J, Lesbros C and Marcelle C (2004) In ovo electroporation of avian somites. Developmental Dynamics 229 (3): 643–650.

Schoenwolf GC (1991) Cell movements driving neurulation in avian embryos. Development 2: 157–168.

Sharma N, Berbari NF and Yoder BK (2008) Ciliary dysfunction in developmental abnormalities and diseases. Current Topics in Developmental Biology 85: 371–427.

Smith CA, Roeszler KN, Ohnesorg T, et al. (2009) The avian Z‐linked gene DMRT1 is required for male sex determination in the chicken. Nature 461 (7261): 267–271.

Song J, McColl J, Camp E, et al. (2014) Smad1 transcription factor integrates BMP2 and Wnt3a signals in migrating cardiac progenitor cells. Proceedings of the National Academy of Sciences of the United States of America 111 (20): 7337–7342.

Stern CD (2005) The chick; a great model system becomes even greater. Developmental Cell 8 (1): 9–17.

Streit A, Tambalo M, Chen J, et al. (2013) Experimental approaches for gene regulatory network construction: the chick as a model system. Genesis 51 (5): 296–310.

Sweetman D, Goljanek K, Rathjen T, et al. (2008) Specific requirements of MRFs for the expression of muscle specific microRNAs, miR‐1, miR‐206 and miR‐133. Developmental Biology 321 (2): 491–499.

Therapontos C, Erskine L, Gardner ER, Figg WD and Vargesson N (2009) Thalidomide induces limb defects by preventing angiogenic outgrowth during early limb formation. Proceedings of the National Academy of Sciences of the United States of America 106 (21): 8573–8578.

Tickle C (1995) Vertebrate limb development. Current Opinion in Genetics & Development 5 (4): 478–484.

Tickle C (2004) The contribution of chicken embryology to the understanding of vertebrate limb development. Mechanisms of Development 121 (9): 1019–1029.

Towers M, Mahood R, Yin Y and Tickle C (2008) Integration of growth and specification in chick wing digit‐patterning. Nature 452 (7189): 882–886.

Voiculescu O, Bertocchini F, Wolpert L, Keller RE and Stern CD (2007) The amniote primitive streak is defined by epithelial cell intercalation before gastrulation. Nature 449 (7165): 1049–1052.

Voiculescu O, Papanayotou C and Stern CD (2008) Spatially and temporally controlled electroporation of early chick embryos. Nature Protocols 3 (3): 419–426.

Wallis JW, Aerts J, Groenen MA, et al. (2004) A physical map of the chicken genome. Nature 432 (7018): 761–764.

Watanabe T, Saito D, Tanabe K, et al. (2007) Tet‐on inducible system combined with in ovo electroporation dissects multiple roles of genes in somitogenesis of chicken embryos. Developmental Biology 305 (2): 625–636.

Yamada T, Placzek M, Tanaka H, Dodd J and Jessell TM (1991) Control of cell pattern in the developing nervous system: polarizing activity of the floor plate and notochord. Cell 64 (3): 635–647.

Yang X, Dormann D, Münsterberg AE and Weijer CJ (2002) Cell movement patterns during gastrulation in the chick are controlled by positive and negative chemotaxis mediated by FGF4 and FGF8. Developmental Cell 3 (3): 425–437.

Yue Q, Wagstaff L, Yang X, Weijer C and Münsterberg A (2008) Wnt3a‐mediated chemorepulsion controls movement patterns of cardiac progenitors and requires RhoA function. Development 135 (6): 1029–1037.

Zamir EA, Czirok A, Cui C, Little CD and Rongish BJ (2006) Mesodermal cell displacements during avian gastrulation are due to both individual cell‐autonomous and convective tissue movements. Proceedings of the National Academy of Sciences of the United States of America 103 (52): 19806–19811.

Zamir EA, Rongish BJ and Little CD (2008) The ECM moves during primitive streak formation – computation of ECM versus cellular motion. PLoS Biology 6 (10): e247.

Further Reading

Brown WR, Hubbard SJ, Tickle C and Wilson SA (2003) The chicken as a model for large‐scale analysis of vertebrate gene function. Nature Reviews Genetics 4 (2): 87–98.

Davey MG and Tickle C (2007) The chicken as a model for embryonic development. Cytogenetic and Genome Research 117 (1–4): 231–239.

Hilgers V, Pourquie O and Dubrulle J (2005) In vivo analysis of mRNA stability using the Tet‐Off system in the chicken embryo. Developmental Biology 284 (2): 292–300.

Hutson MR and Kirby ML (2007) Model systems for the study of heart development and disease. Cardiac neural crest and conotruncal malformations. Seminars in Cell & Developmental Biology 18 (1): 101–110.

Kain KH, Miller JW, Jones‐Paris CR, et al. (2014) The chick embryo as an expanding experimental model for cancer and cardiovascular research. Developmental Dynamics 243 (2): 216–228.

Ogura T (2002) In vivo electroporation: a new frontier for gene delivery and embryology. Differentiation 70 (4–5): 163–171.

Rashidi H and Sottile V (2009) The chick embryo: hatching a model for contemporary biomedical research. Bioessays 31 (4): 459–465.

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Mok, Gi F, Alrefaei, Abdulmajeed F, McColl, James, Grocott, Tim, and Münsterberg, Andrea(Jan 2015) Chicken as a Developmental Model. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0021543]