Chicken Development

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Chicken embryo (day 12)
The chicken (taxon -Gallus gallus) embryo develops and hatches in 20 to 21 days and has been extensively used in embryology studies. Historically, the chicken embryo was one of the first embryos studied, readily available and easy to incubate, embryo development can be directly observed by cutting a small window in the egg shell. A key to this model organism study was the establishment of a staging atlas by Hamburger & Hamilton in 1951 [1], which allowed specifc developmental landmarks to be seen and correlated with experimental manipulations of development. This much cited paper included images of all key stages and was more recently republished in the journal Developmental Dynamics (1993), for a new generation of avian researchers. Probably just as important has been the recent chicken genome sequencing, providing a resource to extend our knowledge of this excellent developmental model.

Fertilized eggs can be easily maintained in humidified incubators and during early stages of development the embryo floats on to of the egg yolk that it is using for nutrition. As the embryo grows it sinks into, or below the, yolk. The regular appearance of somites allowed early experimenters to acurately stage the embryo. The embryo was accessible and easy to manipulate (limb grafts/removal etc) that were informative about developmental processes. Chicken cells and tissues (neural ganglia/fragments) are also easy to grow in tissue culture. The discovery that quail cells have a different nuclear appearance meant that transplanted cells (chick/quail chimeras) could be tracked during development. For example, LeDourian's studies showed how neural crest cells migrate widely throughout the embryo.

This collapsible and sortable table compares the chicken incubation period with other bird species.
Avian Incubation Periods  
Bird Days
Budgerigar 18
Chicken 21
Duck 28
Finch 14
Goose 28
Guinea fowl 28
Muscovy duck 35
Parrot 26
Pheasant 24
Pigeon 18
Quail 16
Swan 35
Turkey 28

Chicken Links: Introduction | Chicken stages | Hamburger Hamilton Stages | Witschi Stages | Placodes | Category:Chicken
Historic Chicken Embryology  
1883 History of the Chick | 1900 Chicken Embryo Development Plates | 1904 X-Ray Effects | 1910 Somites | 1920 Chick Early Embryology | 1933 Neural | 1948 Limb | Movie 1961 | Historic Papers

Chicken Stages

Chicken stages - Hamburger & Hamilton staged the chicken embryo in 1951. The Hamburger Hamilton Stages are most commonly used series for chicken staging. The original paper had approx 25 citations between 1955 - 59, while in the year 1991 alone there were over 300 citations. Series of Embryonic Chicken Growth. J. Morphology, 88 49 - 92 (1951). Atlas recently republished by J.R. Sanes in Developmental Dynamics 195 229-275 (1993).

Hamburger Hamilton Stages (1951)  
Hamburger Hamilton Stages
Identification of Stages   (Chicken Development)
Before Laying
Early cleavage
3.5-4.5 hr Shell membrane of egg formed in isthmus of oviduct
During cleavage
Germ wall formed from marginal periblast
Late cleavage
4.5-24.0 hr Shell of egg formed in uterus
After Laying
Preprimitive streak (embryonic shield)
6-7 hr Initial primitive streak, 0.3-0.5 mm long
12-13 hr Intermediate primitive streak
18-19 hr Definitive primitive streak, ±1.88 mm long
19-22 hr Head process (notochord)
23-25 hr Head fold
23-26 hr 1 somite; neural folds
7 to 8-
ca. 23-26 hr 1-3 somites; coelom
26-29 hr 4 somites; blood islands
29-33 hr 7 somites; primary optic vesicles
9+ to 10-
ca. 33 hr 8-9 somites; anterior amniotic fold
33-38 hr 10 somites; 3 primary brain vesicles
40-45 hr 13 somites; 5 neuromeres of hindbrain
45-49 hr 16 somites; telencephalon
48-52 hr 19 somites; atrioventricular canal
13+ to 14-
ca. 50-52 hr 20-21 somites; tail bud
50-53 hr 22 somites; trunk flexure; visceral arches I and II, clefts 1 and 2
14+ to 15-
ca. 50-54 hr 23 somites; premandibular head cavities
50-55 hr 24-27 somites; visceral arch III, cleft 3
51-56 hr 26-28 somites; wing bud; posterior amniotic fold
52-64 hr 29-32 somites; leg bud; epiphysis
3 da 30-36 somites extending beyond level of leg bud; allantois
3.0-3.5 da 37- 40 somites extending into tail; maxillary process
3.0-3.5 da 40-43 somites; rotation completed; eye pigment
3.5 da 43-44 somites; visceral arch IV, cleft 4
3.5-4.0 da Somites extend to tip of tail
4 da Dorsal contour from hindbrain to tail is a curved line
4.5 da Toe plate
4.5-5.0 da Elbow and knee joints
5 da 1st 3 toes
5.0-5.5 da Beak
5.5-6.0 da 3 digits, 4 toes
6.0-6.5 da Rudiment of 5th toe
6.5-7.0 da Feather germs; scleral papillae; egg tooth
7.0-7.5 da Web between 1st and 2nd digits
7.5 da Anterior tip of mandible has reached beak
7.5-8.0 da Web on radial margin of wing and 1st digit
8 da Nictitating membrane
8.5-9.0 da Phalanges in toes
10 da Length of 3rd toe from tip to middle of metatarsal joint = 5.4 ±0.3 mm; length of beak from anterior angle of nostril to tip of bill = 2.5mm; primordium of comb; labial groove; uropygial gland
11 da Length of 3rd toe = 7.4 ±0.3mm; length of beak = 3.0 mm
12 da Length of 3rd toe = 8.4 ± 0.3 mm; length of beak = 3.1 mm
13 da Length of 3rd toe = 9.8 ± 0.3 mm; length of beak = 3.5 mm
14 da Length of beak = 4.0 mm; length of 3rd toe = 12.7 ± 0.5 mm
15 da Length of beak from anterior angle of nostril to tip of upper bill = 4.5 mm; length of 3rd toe = 14.9 ± 0.8 mm
16 da Length of beak = 4.8 mm; length of 3rd toe = 16.7 ± 0.8 mm
17 da Length of beak = 5.0 mm; length of 3rd toe = 18.6 ± 0.8 mm
18 da Length of beak = 5.7 mm; length of 3rd toe = 20.4 ± 0.8 mm
19-20 da Yolk sac half enclosed in body cavity; chorio-allantoic membrane contains less blood and is "sticky" in living embryo
20-21 da Newly-hatched chick

Hamburger V. and Hamilton HL. A series of normal stages in the development of the chick embryo. (1951) J Morphol. 88(1): 49-92. PMID 24539719 PDF

Original 1951 paper (and all data) was republished in 1992. V Hamburger, H L Hamilton A series of normal stages in the development of the chick embryo. 1951. Dev. Dyn.: 1992, 195(4);231-72 PubMed 1304821


Note that there was also an earlier Witschi staging, and a 1900 staging series by Franz Keibel and Karl Abraham[2], and an earlier (1883) series by Foster, Balfour, Sedgwick, and Heape.[3]

Normal Plates of the Development of the Chicken Embryo (1900)

Links: Chicken Stages | Hamburger Hamilton | Witschi | 1900 | 1883 | PDF Poster- Hamburger Hamilton Stages | 2006 reproduction of the original paper

Some Recent Findings

  • FGF8 coordinates tissue elongation and cell epithelialization during early kidney tubulogenesis[4] "When a tubular structure forms during early embryogenesis, tubular elongation and lumen formation (epithelialization) proceed simultaneously in a spatiotemporally coordinated manner. We here demonstrate, using the Wolffian duct (WD) of early chicken embryos, that this coordination is regulated by the expression of FGF8, which shifts posteriorly during body axis elongation. FGF8 acts as a chemoattractant on the leader cells of the elongating WD and prevents them from epithelialization, whereas static ('rear') cells that receive progressively less FGF8 undergo epithelialization to form a lumen. Thus, FGF8 acts as a binary switch that distinguishes tubular elongation from lumen formation. The posteriorly shifting FGF8 is also known to regulate somite segmentation, suggesting that multiple types of tissue morphogenesis are coordinately regulated by macroscopic changes in body growth." Fibroblast Growth Factor
  • 4D fluorescent imaging of embryonic quail development[5] "Traditionally, our understanding of developmental biology has been based on the fixation and study of embryonic samples. Detailed microscopic scrutiny of static specimens at varying ages allowed for anatomical assessment of tissue development. The advent of confocal and two-photon excitation (2PE) microscopy enables researchers to acquire volumetric images in three dimensions (x, y, and z) plus time (t). Here, we present techniques for acquisition and analysis of three-dimensional (3D) time-lapse data. Both confocal microscopy and 2PE microscopy techniques are used. Data processing for tiled image stitching and time-lapse analysis is also discussed. The development of a transgenic Japanese quail system, as discussed here, has provided an embryonic model that is more easily accessible than mammalian models and more efficient to breed than the classic avian model, the chicken."
More recent papers  
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This table shows an automated computer PubMed search using the listed sub-heading term.

  • Therefore the list of references do not reflect any editorial selection of material based on content or relevance.
  • References appear in this list based upon the date of the actual page viewing.

References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

Links: References | Discussion Page | Pubmed Most Recent | Journal Searches

Search term: Chicken Embryology

Yalin Cheng, Bin Gao, Haitao Wang, Naijian Han, Shimiao Shao, Shaoyuan Wu, Gang Song, Yong E Zhang, Xiaojia Zhu, Xin Lu, Yanhua Qu, Fumin Lei Evolution of beak morphology in the Ground Tit revealed by comparative transcriptomics. Front. Zool.: 2017, 14;58 PubMed 29299037

B U Metzler-Zebeli, E Magowan, M Hollmann, M E E Ball, A Molnár, K Witter, R Ertl, R J Hawken, P G Lawlor, N E O'Connell, J Aschenbach, Q Zebeli Differences in intestinal size, structure, and function contributing to feed efficiency in broiler chickens reared at geographically distant locations. Poult. Sci.: 2017; PubMed 29253222

Susann Brosig, N Peukert, R Metzger, H Schneider, R Haak, J Gosemann, M Lacher, D Kluth X-ray micro-computed-tomography in pediatric surgery: a new tool for studying embryos. Pediatr. Surg. Int.: 2017; PubMed 29167984

Guangwei Hu, Guang Li, Hui Wang, Yiquan Wang Hh gene participates in the left-right asymmetry development of amphioxus by controlling Cer expression. Development: 2017; PubMed 29122841

Pieter Vancamp, Marie-Anne Deprez, Michiel Remmerie, Veerle M Darras Deficiency of the thyroid hormone transporter MCT8 in neural progenitors impairs cellular processes crucial for early corticogenesis. J. Neurosci.: 2017; PubMed 29109240

Gallus gallus

Taxonomy Id: 9031

Preferred common name: chicken

Rank: species

Genetic code: Translation table 1 (Standard) Mitochondrial genetic code: Translation table 2

Other names: dwarf Leghorn chickens (includes), red jungle fowl (includes), chickens (common name), Gallus domestics (misnomer), Gallus galls domesticus (misnomer)

Lineage (abbreviated ): Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Archosauria; Aves; Neognathae; Galliformes; Phasianidae; Phasianinae; Gallus

Chicken Movies

Chicken movie 1961.jpg
 ‎‎Chicken (1961)
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Mesoderm migration movie 1 icon.jpg
 ‎‎Mesoderm Move
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Chicken Embryo Somite1-icon.jpg
 ‎‎Chicken Somite
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Chick Heart 001-icon.jpg
 ‎‎Normal Heart
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Chick Heart 002-icon.jpg
 ‎‎Abnormal Heart 1
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Chick Heart 002-icon.jpg
 ‎‎Abnormal Heart 2
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 ‎‎GIT Motility
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Neural crest migration Chicken Head (movies overview)
 ‎‎Neural Crest 1
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 ‎‎Neural Crest 2
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 ‎‎Neural Crest 3
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 ‎‎Neural Crest 4
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 ‎‎Neural Crest 5
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 ‎‎Neural Crest 6
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 ‎‎Neural Crest 7
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Chicken Placode
Chicken Placode Movie 6 icon.jpg
 ‎‎Placode 6
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Chicken Placode Movie 1 icon.jpg
 ‎‎Placode 1
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Chicken Placode Movie 2 icon.jpg
 ‎‎Placode 2
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Chicken Placode Movie 3 icon.jpg
 ‎‎Placode 3
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Chicken Placode Movie 4 icon.jpg
 ‎‎Placode 4
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Chicken Placode Movie 5 icon.jpg
 ‎‎Placode 5
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Chicken Placode Movie 7 icon.jpg
 ‎‎Placode 7
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Chicken Placode Movie 8 icon.jpg
 ‎‎Placode 8
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Chicken Placode Movie 9 icon.jpg
 ‎‎Placode 9
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Links: Movies

Other Chicken Atlases

Vertebrate and Invertebrate Embryos (7th Edition) G.C. Schoenwolf, Prentice Hall, New Jersey

An Atlas of Embryology (1975) W.H. Freeman and B. Bracegirdle, Heinemann Educational Books, UK.

This is an ATLAS (no description of development) , basically reprinted from the original 1963 edition.

Photos with labelled diagrams covering Amphioxus (worm) Frog, Chicken.

An Atlas for Staging Mammalian and Chick Embryos (1987) H. Bultler and B.H. Juurlink, CRC Press Inc., Florida

This ATLAS is not a complete series of development but has interesting comparisons of species.

Mostly photos of embryos with a few drawn diagrams and a series of staging correlation graphs.

Bird Evolution

Birds and Dinosaurs? as quoted in a Curent Biology review "...abundant and ever increasing evidence places birds as one surviving lineage of the diverse clade Dinosauria"[6][7]

Chicken Genomics

The first draft of the chicken genome was publicly released in March, 2004. There are a number of sites that have begun looking into establishing chicken genomics partly due to its powerful history as a model of vertebrate development that is easy to observe, manipulate and is also cheap. (see also NIH Proposal for Chicken Genomics | NCBI Chicken Genome Resources)

A summary of chicken genome resources has recently been identified in a review in Developmental Dynamics by Antin PB and Konieczka JH.[8]

Chicken Sex Determination

In chicken development sex determination depends on a ZZ male/ZW female mechanism.

This differs from mammalian sex determination which is based upon testis expression of an Sry gene in somatic supporting Sertoli cells.

In the gonad, the coelomic epithelium contributes only to non-steroidogenic interstitial cells and nephrogenous mesenchyme contributes both Sertoli cells and steroidogenic cells.


Chicken primordial germ cell migration model.jpg

Primordial Germ Cell Migration Model[9]

HH12–13 - yolk sac circulation courses in loop (red arrows) to enter the embryo via the heart. The majority of PGCs (green dots) localized axially at the border between the area opaca and pellucida, where the sinus terminalis converged in the anterior vitelline veins. HH14–16 - PGCs (green dots) circulated effectively towards the embryo via the sinus terminalis and the anterior vitelline veins towards the heart. Then PGCs traffic via the aorta to the caudal part of the embryo and become lodged in the genital ridges.

Chicken Heart

Note these are Hamburger Hamilton Stages of chicken development, see also Heart 3D reconstruction.

Chicken Cardiac Stages

From review [10]

  • HH 8 (26–29 hours, 4–6 somites)
  • HH 9 (29–33 hours, 7–9 somites) - Cardiac neural crest cells begin the process of EMT and emigrate from the neural tube.
  • HH 10–11 (33–45 hours, 10–15 somites) - Primary heart tube
  • HH 12-13- (45–49 hours, 16–19 somites) - dextral-looping phase of looping completed at stage 12.
  • HH 13+ (50–52 hours, 20–21 somites) - c-shaped heart loop transformed into the s-shaped heart loop. Cardiac neural crest has stopped producing cells.

Chicken Somitogenesis

Somitogenesis 01 icon.jpg
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Chicken Embryo Somite1-icon.jpg
 ‎‎Chicken Somite
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Gene expression Somite timing


Chick somitogenesis oscillator[11]

Chicken body elongation model.jpg

Chicken body elongation model[12]

Chicken Limb

Limb hairy2 expression model.jpg

Limb Hairy2 Expression Model[13]

Hairy2 is a "molecular oscillator" involved in both somite and limb development.

Chicken limb gene expression 03.jpg

Chicken stage 21 to 27 wing bud Tbx2 and Tbx3 expression[14]


The following gene expression data is from a study of different head regions during development.[15]

  • Frontonasal Prominence - CASH1/ASCL1, POSTN, OGN, CYP26A1, NR2E1, and SCARA5.
    • Olfactory epithelium SP8, EYA2, and SIX3
  • Maxillary/Trigeminal Ganglion - SOX10, TAGLN3.
  • Mandibular - DLX1, HAND2 (highest), LHX8, MSX2, PITX2, and TWIST2.
    • Mandibular/maxillary prominences differentially expressed - BETA3, HAND2, and MSX2.

Historic Studies

The Elements of Embryology - Volume 1 by Foster, M., Balfour, F. M., Sedgwick, A., & Heape, W. (1883)

The History of the Chick: Egg structure and incubation beginning | Summary whole incubation | First day | Second day - first half | Second day - second half | Third day | Fourth day | Fifth day | Sixth day to incubation end

Elements of Embryology - Volume 1 - Figures


  1. V Hamburger, H L Hamilton A series of normal stages in the development of the chick embryo. 1951. Dev. Dyn.: 1992, 195(4);231-72 PubMed 1304821
  2. Keibel F. and Abraham K. Normal Plates of the Development of the Chicken Embryo (Gallus domesticus). (1900) Vol. 2 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
  3. Foster M. Balfour FM. Sedgwick A. and Heape W. The Elements of Embryology (1883) Vol. 1. (2nd ed.). London: Macmillan and Co.
  4. Yuji Atsuta, Yoshiko Takahashi FGF8 coordinates tissue elongation and cell epithelialization during early kidney tubulogenesis. Development: 2015, 142(13);2329-37 PubMed 26130757
  5. Christie A Canaria, Rusty Lansford 4D fluorescent imaging of embryonic quail development. Cold Spring Harb Protoc: 2011, 2011(11);1291-4 PubMed 22046043
  6. Julia Clarke, Kevin Middleton Bird evolution. Curr. Biol.: 2006, 16(10);R350-4 PubMed 16713939
  7. Bent E K Lindow, Gareth J Dyke Bird evolution in the Eocene: climate change in Europe and a Danish fossil fauna. Biol Rev Camb Philos Soc: 2006, 81(4);483-99 PubMed 16893476
  8. Parker B Antin, Jay H Konieczka Genomic resources for chicken. Dev. Dyn.: 2005, 232(4);877-82 PubMed 15739221 | Developmental Dynamics
  9. Ana De Melo Bernardo, Kaylee Sprenkels, Gabriela Rodrigues, Toshiaki Noce, Susana M Chuva De Sousa Lopes Chicken primordial germ cells use the anterior vitelline veins to enter the embryonic circulation. Biol Open: 2012, 1(11);1146-52 PubMed 23213395 | PMC3507194 | Biol Open
  10. Brad J Martinsen Reference guide to the stages of chick heart embryology. Dev. Dyn.: 2005, 233(4);1217-37 PubMed 15986452
  11. Gennady Tenin, David Wright, Zoltan Ferjentsik, Robert Bone, Michael J McGrew, Miguel Maroto The chick somitogenesis oscillator is arrested before all paraxial mesoderm is segmented into somites. BMC Dev. Biol.: 2010, 10;24 PubMed 20184730
  12. Isabel Olivera-Martinez, Hidekiyo Harada, Pamela A Halley, Kate G Storey Loss of FGF-dependent mesoderm identity and rise of endogenous retinoid signalling determine cessation of body axis elongation. PLoS Biol.: 2012, 10(10);e1001415 PubMed 23118616 | PLoS Biol.
  13. Caroline J Sheeba, Raquel P Andrade, Isabel Palmeirim Joint interpretation of AER/FGF and ZPA/SHH over time and space underlies hairy2 expression in the chick limb. Biol Open: 2012, 1(11);1102-10 PubMed 23213390 | PMC3507187 | Biol Open
  14. Malcolm Fisher, Helen Downie, Monique C M Welten, Irene Delgado, Andrew Bain, Thorsten Planzer, Adrian Sherman, Helen Sang, Cheryll Tickle Comparative analysis of 3D expression patterns of transcription factor genes and digit fate maps in the developing chick wing. PLoS ONE: 2011, 6(4);e18661 PubMed 21526123 | PLoS One.
  15. Marcela Buchtová, Winston Patrick Kuo, Suresh Nimmagadda, Shari L Benson, Poongodi Geetha-Loganathan, Cairine Logan, Timothy Au-Yeung, Eric Chiang, Katherine Fu, Joy M Richman Whole genome microarray analysis of chicken embryo facial prominences. Dev. Dyn.: 2010, 239(2);574-91 PubMed 19941351



Matthew J Korn, Karina S Cramer Windowing chicken eggs for developmental studies. J Vis Exp: 2007, (8);306 PubMed 18989413

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